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Status Survey and Conservation Action Plan

Cactus and Succulent Plants

Compiled by Sara Oldfield

IUCN/SSC Cactus and Succulent Specialist Group

Sultanate of Oman

The World Consewation Union SI'LCI~S Suitv1v.4~ COMMISSI~)N

m @ Chicago Zoological Society

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Oldfield, Sara (camp.) (1997). Cactus and Succulent Plants- Status Sun/ey and Consen/ation Action Plan. IUCN/SSC Cactus and Succulent Specialist Group. IUCN, Gland, Switzerland and Cambridge, UK. IO + 212 pp.

2-8317-0390-5

Pachypodium namaquanum, the renowned ‘Half Mens’ from the Richtersveld, at one time listed on CITES Appendix I; now considered relatively safe, but still Vulnerable in parts of its range. (National Botanical Institute)

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Contents

Foreword ......................................... V

Acknowledgements ................................ vi

Executive Summary ............................... vii

(English, Spanish, French)

Acronyms ........................................ X

Introduction ...................................... 1

Chapter 1. Taxonomic Groups ..................... .3 Agavaceae .................................. ...3 Aizoaceae .................................. ...8 Aloaceae.....................................lO Asclepiadaceae. ............................... 14 Cactaceae.. ................................ ..17 Crassulaceae..................................2 0 Euphorbiaceae ............................... .23 Portulacaceae .............................. ..2 6

Chapter 2. Conservation Measures ................. .28 National legislation ........................... .28 International legislation ....................... .30

The Convention on Biological Diversity. ....... 30 The Berne Convention ..................... .30 The EC Habitats Directive. ................. .30

Controlling the trade. ......................... .31 CITES....................................3 1 Illegal trade. .............................. .36 Trade in non-CITES succulents. ............. .37

In situ conservation ........................... .37 Protecting the habitats of threatened species. .. .37 Protected areas ........................... .38 International designations .................. .38 National designations ...................... .40 Private land. .............................. .41 Species management. ...................... .42

Ex situ conservation. .......................... .42 Botanic garden networks ................... .43 Seedbanks.. ............................ ..4 5 Botanic gardens ........................... .45 Hobbyist collections and nurseries ........... .49

Chapter 3. Regional Accounts. .................... .52 Tropical Africa. .............................. .52

Somalia-Masai regional centre of endemism ... .53 Kenya .................................. ..5 3 Ethiopia ................................. .55 Somalia. ................................. .57

Madagascar...................................5 9 Threats ................................. ..6 1 Conservation status. ....................... .63 Priority sites for conservation. ................ 64 Existing conservation measures ............... 65

Southern Africa .............................. .67 Distribution in southern Africa ............... 69 Conservation status. ....................... .70 Threats ................................. ..7 2 Existing conservation measures ............... 75 Proposed conservation strategy .............. .85

North America. .............................. .85 Succulent plant flora in the United States ...... 86 Threats ................................. ..8 6 Local uses and commercialisation ............ .86 Existing conservation measures .............. .87 Assessment of existing conservation measures. . .88

Mexico.......................................8 9 Succulent plant flora. ...................... .89 Localuse.................................. 0 Threats ................................. ..9 0 Conservation status. ....................... .93 Priority sites for succulent plant conservation .. .94 Existing conservation measures ............... 95

The West Indies. ............................. .99 The succulent flora: plant communities and

sites of occurrence ....................... 100 Threats.. .............................. ..lO 3 Conservation status. ....................... 106 Priority sites for conservation. ............... 106 Existing conservation measures .............. 109

South America ............................... 111 The Caribbean Region ..................... 111 The Andean Region ....................... 112 The Brazilian Region ...................... 118 The Chile-Patagonian Region ............... 125

Chapter 4. Action Proposals ...................... .128

General.....................................12 8 Ex situ conservation. .......................... 129 Taxon specific. ............................... 130

Agavaceae................................13 0 Aloaceae.................................13 1 Crassulaceae. ............................. 131

Regional action proposals. ..................... 131 Canary Islands ............................ 131 Mediterranean region. ..................... 131 Somalia .................................. 131 Southern Africa ........................... 132 Madagascar ............................ ..13 5 India .................................. ..13 6 United States of America. .................. 136 Mexico and United States of America ........ 138 Mexico...................................13 8 Central America .......................... 140 Caribbean. ............................... 140 South America. ........................... 142 Columbia/Venezuela. ...................... 142 Venezuela................................14 3

Ecuador..................................14 3 Peru.....................................14 3 Chile .................................. ..14 3 Brazil....................................14 3 Paraguay ................................. 144

References ...................................... 145

Annex 1 Agavaceae with restricted distribution ..... 156

Annex 2 Asclepiadaceae of conservation concern ... 159

Annex 3 Succulents regulated by CITES . . . . . . . .

Annex 4 Kenyan succulents of highest conservation priority. . . . . . . . . . . . . . . .

Annex 5 Succulents of the Mediterranean Region

Annex 6 Succulents of the Canary Islands. . . . . . .

Annex 7 Succulents of Madagascar. . . . . . . . . . . . .

. . 164

. . 165

. . 166

I . 171

, . 174

Annex 8 Threatened succulents recorded for the Flora of southern Africa (FSA) region . . . 179

Annex 9 Threatened succulents of Zimbabwe. . . . . . . 185

Annex 10 Threatened succulents of India . . . . . . . . . . . 186

Annex 11 Threatened succulents of Mexico . . . . . . . . . 189

Annex 12 Mexican succulent nurseries. . . . . . . . . . . . . . 191

Annex 13 Mexican botanic gardens . . . . . . . . . . . . . . . . 192

Annex 14 Succulents of the West Indies . . . . . . . . . . . . 194

Annex 15 Brazilian cacti. . . . . . . . . . . . . . . . . . . . . . . . . . 199

Annex 16 The IUCN Red List Categories. . . . . . . . . . .203

Annex 17 IUCN/SSC Cactus and Succulent Plant Specialist Group members and contributing authors. . . . . . . . . . . . . . . . . . .208

iv

Foreword

The sight was depressing. We looked closely at the huge pile of dead cacti to see if any had survived, but none were found. Commercial collectors had dug up these hundreds of plants, but for some reason had simply left them behind to die. I had seen the destruction of cactus populations in other places, where whole hillsides of cacti had been removed and carried away, but nowhere nearly as many plants had been ruthlessly removed and left to die as in this remote area of the Chihuahuan Desert. Devastation of populations of cacti and succulents has occurred in many parts of the world, sometimes through greed and selfishness of people who wish to make money from the plants, but at other times as land is converted to agriculture, bulldozed for houses or highways, or flooded to create dams.

For 40 years I have travelled throughout the arid regions of both North and South America doing research on cacti. I have been fortunate to see some of the rarest cacti known, but have also witnessed their pillage by people who fail to understand their importance in the wild and who do not care if they are destroyed, thus depriving future generations of the experience of seeing them. The same things are happening in areas of Africa and Madagascar. Cacti and succulents face terrible threats as the human population grows and land is converted from its natural state. Scientists are struggling to learn what they can before some of these populations of plants disappear, but it is discouraging to see tagged plants removed from study sites by unscrupulous collectors, or a study area bulldozed to make a wider road. At times the frustration has been so great that one is led to wonder if conservation activities are worth the time and effort. But then one hears the statement that “if only people knew what was happening, then they might stop this needless destruction.” Or we see a piece of countryside that has been set aside specifically to protect and preserve a population of rare plants. We may read a report that collectors have been arrested for illegally removing plants. These are what give us hope; perhaps we can inform and educate people of the importance of preserving cacti and succulents in the wild, and, as a result, species can be saved from extinction.

This Action Plan is a much-needed publication on cacti and succulents, for it brings together data that have never been compiled before, which should help greatly in conservation efforts of cacti and succulents throughout the world. Included is information on eight succulent families, as well as nine geographic regions of the world, plus conservation strategies including trade control. However, the most important part of the Action Plan is the section dealing with proposals for conservation actions. These are both timely and practical, for experts from various regions have described how conservation efforts can be carried out. It is hoped that these proposals will stimulate governments, foundations, other organisations and individuals to respond to this ever- increasing crisis concerning the conservation of cacti and succulents.

It has been my privilege to serve as Chair of the Cactus and Succulent Specialist Group of the Species Survival Commission for nearly ten years. The conception and writing of this Action Plan is clearly the most significant contribution of this Group. It has been a challenge developing this work, for the Group wanted the expertise of the world’s authorities, who are often heavily committed to other projects. Those that responded with contributions have made this Action Plan unique and of considerable value. Their willingness to contribute chapters or sections is greatly appreciated. Special thanks go to Sara Oldfield, Secretary of the Group, for overseeing and compiling the material as it was submitted. The project would never have been possible without her efforts. Wendy Strahm and Robin Sears of the Species Survival Commission also deserve many thanks for seeing the Plan through to its final production.

As one looks at this Action Plan, I ask that it be read with deep appreciation, not only of the plants described, but also for the contributions of so many who work with them. Hopefully, this publication will facilitate conservation efforts on behalf of the cacti and succulents, a remarkable group of plants that must be preserved for eternity.

Dr. Edward F. Anderson Chair, IUCN/SSC Cactus and Succulent Specialist Group

Acknowledgements

The preparation of this SSC Action Plan has been a collaborative and largely voluntary venture to which many specialists have given freely of their time and expertise. Authors and contributors to the Plan are listed in Annex 17. Their work on this document, both in the initial preparation of the manuscripts, and subsequent help with additional information, review, and provision of illustrations is greatly appreciated. In turn, the authors have consulted widely in the preparation of their accounts and acknowledgements are given at the end of individual sections as appropriate. All members of the SSC Group have been involved in the preparation of the SSC Action Plan in some way, together with many members of the 10s and experts in conservation organisations. Sincere thanks to everyone involved. Special thanks are due to Ted Anderson, Susan Carter-Holmes, Craig Hilton- Taylor, Gideon Smith, Diedrich Supthut, and Nigel Taylor for guidance and encouragement in developing the

Action Plan, and to Wendy Strahm and Robin Sears for ensuring its completion. Robin Sears played a particularly important role in assisting with the final editing, picture research and preparation of the report for publication. Thanks also go to Gary Lyons and Rob Wallace for final review of the text; and to Juan Manuel Lopez Ramirez for the Spanish translation, and Paul Strahm and Bertrand de Montmollin for the French translation of the Executive Summary. Institutional support is acknowledged from the Desert Botanical Garden, Phoenix; Royal Botanic Gardens, Kew; and World Conservation Monitoring Centre. Financial support for preparation of the Action Plan was provided by the Peter Scott IUCN/SSC Action Plan fund.

Sara Oldfield Secretary, IUCN/SSC Cactus and Succulent Specialist Group

vi

Executive Summary

The often bizarre growth-forms and attractive flowers of cacti and other succulents have promoted widespread interest in this group of plants and horticultural popularity worldwide. Succulent plants are also of great ecological and economic significance, particularly in arid and semi-arid parts of the world. Although the definition of succulence as applied to plants is constantly under debate, about 10,000 plant species are generally recognised as succulent, within thirty plant families.

Of these succulent plant species, an estimated 2000 species are threatened with global extinction in the wild, and many more are regionally or nationally threatened. Habitat destruction is the major threat, and in common with other horticulturally desirable plant groups, overcollection for international trade remains a significant problem. The Cactus and Succulent Plants Action Plan, produced by members of the Species Survival Commission of IUCN-The World Conservation Union, brings together current information, never before compiled, on the population status, threats, and conservation of this group of important plants from around the world. From this compilation, priorities for conservation action are emphasised, providing direction for funding in plant conservation work. Conservationists, scientists, government officials, protected area managers, educators, and grant awarding bodies alike should find this document helpful in their work to conserve global and local flora. The contributors to this Action Plan encourage collaborative work among these interested parties.

The publication comprises four chapters and a series of annexes that provide readers with concise information on the current status of cactus and succulent populations. The extensive bibliography provides a comprehensive resource for more information on this group of plants. The Plan begins with overviews, written by botanists who specialise in the study of these particular plant families, of the distribution, diversity, threats, and status of eight main taxonomic groups of succulents including the Agavaceae, Aizoaceae, Aloaceae, Asclepiadaceae, Cactaceae, Crassulaceae, Euphorbiaceae, and Portulacaceae. Some of these groups are of great economic importance, many in the ornamental trade industry, and others, such as the Agavaceae, in the fibre and food industries.

Chapter 2 of the Action Plan describes and reviews existing conservation measures for succulent plants around the world with information on legislation, controlling the trade, and in situ and ex situ conservation. The intention of this chapter is to identify successful conservation activities which can be used as models elsewhere, and also to highlight priorities for further action. It is particularly important in reviewing international conservation measures to show how succulent plant conservation needs can be linked into broader initiatives and frameworks for biodiversity conservation.

Action for succulent plant conservation must take place primarily at the national and local levels and be implemented as far as possible by in-country agencies and local experts. This is accepted throughout the Action Plan, and Chapter 3, the regional accounts, has been largely prepared by experts within the regions concerned. Chapter 3 concentrates on the regions of the world which have the major concentrations of succulent p lants.

The final chapter of the Plan describes the priority conservation action proposals, developed by the members of the SSC Cactus and Succulent Plant Specialist Group, for succulent plants around the world. Implementing these proposals will save the maximum diversity of succulents based on our present knowledge.

The SSC Cactus and Succulent Specialist Group calls for:

Field research to support understanding of the taxonomy and conservation status of succulent plants, Increased in situ protection for succulent plant species through the development of protected area networks, Coordinated ex situ protection of threatened succulent species to support the conservation of species in their natural habitats wherever possible, Effective national legislation for all threatened succulent plant species, Effective trade controls for all wild succulent plant species threatened by exploitation for international commerce, Education on the value of succulent plants and the need for their conservation and sustainable use.

vii

Redmen

Las a menudo extranas formas de crecimiento y las flores atractivas de cactus y otras plantas suculentas han logrado un inter& general por este grupo de plantas y una popularidad para su utilization en horticultura en todo el mundo. La s plantas suculentas importancia ecologica y economica

son tambie n de gran icularm ente en las

zonas aridas y semiaridas de todo el mundo. Aunque la definition de suculencia aplicada a las plantas esta constantemente en debate, cerca de unas 10.000 especies de plantas, clasificadas en unas 30 familias, son reconocidas generalmente

De estas especies de corn0 planta

suculentas. s suculentas, unas 2.000

estan globalmente amenazadas en la naturaleza con la extincion, y muchas otras en peligro regional o national. La destruction de1 habitat es la mayor amenaza, y es compartida con otros grupos atractivos de plantas de jardineria, en donde su sobreexplotacion para el comercio international es un problema de gran trascendencia. El Plan de Action para Cactus y Plantas Suculentas, que elaboraron 10s miembros de la Comision de Supervivencia de Especies (CSE) de la UICN - Union Mundial para la

directrices para el financiamiento de1 trabajo de conservation de las plantas. Conservacionistas, cientificos, funcionarios, gestores de areas protegidas, educadores y organismos financiadores, de1 mismo modo, podrian encontrar ayuda en este document0 para sus trabajos de conservation de la flora local y global. Los contribuidores a este Plan de Action animan al trabajo de colaboracion entre estos grupos interesados.

La publication consta de quatro capitulos y una serie de anexos que proporcionan a 10s lectores una information concisa sobre el estado actual de las poblaciones de cactus y suculentas. La extensa bibliografia proporciona una amplia fuente de information suplementaria sobre cactus y plantas suculentas. El Plan comienza con resumenes, que escriben botanicos especializados en el estudio de estas familias especificas de plantas, acerca de la distribution, diversidad, amenazas y el estado de echo grupos taxonomicos principales de cactus y suculentas entre las que se encuentran las familias Agavaceae, Aizorceae, Aloaceae, Asclepiadaceae, Cactaceae, Crassulaceae, Euphorbiaceae, y Portulaceae. Algunos de estos grupos son de gran importancia economica, muchos de ellos incluidos en el comercio de ornamentales. v otros coma las Agavaceas, en las industrias alimentaria y textil.

El Capitulo 2 de este Plan de Action describe y examina las medidas de conservation existentes para las plantas suculentas de todo el mundo, con information sobre legislation, control de1 comercio y conservation in situ y ex situ. El objetivo de este capitulo es identificar las actividades exitosas de conservation que pueden ser usadas como modelos en otros lugares y muestra ademas las prioridades para una action mas completa. Es particularmente importante en el analisis de las medidas de conservation international, mostrar coma las necesidades de conservation de las plantas suculentas pueden ser unidas dentro de las iniciativas generales de conservation y en 10s marcos para la conservation de la biodiversidad.

La action para la conservation de las plantas suculentas debe tomarse ante todo en 10s niveles locales y nacionales y ser llevada a cabo tanto coma sera possible por agencias estatales y locales especializadas. Esto es aceptado en todo el Plan de Action, y en el Capitulo 3,10s informes regionales han sido preparados por expertos de las propias regiones. El Capitulo 3 se concentra en las regiones de1 mundo que tienen el mayor numero de plantas suculentas.

El capitulo final describe las propuestas de acciones prioritarias para la conservation para las plantas suculentas de todo el mundo. Lo elaboraran miembros de1 Grupo Especialista de la CSE/UICN en Cactaceas y Suculeantas. Hacienda efectivas estas propuestas, basadas en nuestros conocimientos actuales, salvaremos al maxim0 la diversidad de las suculentas.

El Grupo Especialista CSE en Cactus y Suculentas pide:

Una investigation de campo para apoyar el entendimiento de la taxonomia y el estado de conservation de las plantas suculentas, Aumento de la protection in situ de las especies de plantas suculentas a traves de1 desarrollo de redes de areas protegidas, Coordination de la protection ex situ de las plantas suculentas para apoyar la conservation de especies en sus hgbitats naturales dondequiera que sea posible, Una legislation national eficaz para todas las especies de plantas suculentas amenazadas, Controles efectivos sobre el comercio de todas las plantas suculentas silvestres amenazadas por la sobreexplotacion en el comercio international, Education sobre el valor de las plantas suculentas y la necesidad de conservation y uso sostenible.

. . . VIII

R&urn6

L’aspect souvent particulier ainsi que la beaute des fleurs des cactus et d’autres plantes succulentes suscitent beaucoup d’interet de la part du public pour ce type de vegetation dont la culture est tres repandue. Les plantes succulentes ont egalement une grande importance ecologique et economique a travers le monde, particulierement dans les regions arides ou semi-arides. Bien que la definition de ce quest exactement une plante “succulente” fasse toujours l’objet de discussions, on peut considerer que quelque 10,000 especes, subdivisees en trente familles, font partie de ce groupe de planks.

Au niveau mondial, on estime que pres de 2,000 especes de plantes succulentes sont menacees d’extinction dans leur habitat naturel. Un nombre bien plus eleve est menace de disparition a une echelle regionale ou nationale. La menace la plus importante pour les plantes succulentes est la destruction de leurs habitats, puis, comme pour beaucoup d’autres groupes de plantes attractives, une recolte trop intensive pour satisfaire les besoins du commerce international constitue un probleme tout particulier.

Le Plan d’Action pour les cactus et les plantes succulentes publie par les membres de la Commission de la sauvegarde des especes (CSE) de IUnion mondiale pour la nature (UICN) r&nit toutes les informations au niveau mondial - ce qui n’avait encore jamais ete fait - au sujet de la repartition, des menaces et de la conservation de cet important groupe de plantes. Sur la base de cette compilation, des priori& de conservation sont degagees, en indiquant les directions a suivre pour rechercher des financements pour les actions de conservation. Ce document constitue une base de travail pour la conservation de la flore au niveau mondial ou local a l’intention des acteurs de la conservation, des scientifiques, des dirigeants gouvernementaux, des gestionnaires d’espaces proteges, des enseignants et des organes de subventionnement. Les auteurs du Plan d’Action encouragent une collaboration etroite entre toutes les parties concernees.

Cet ouvrage, constitue de quatre chapitres et d’une serie d’annexes, offre au lecteur des renseignements concis sur l’etat actuel de conservation des populations de cactus et de plantes succulentes. Une abondante bibliographie offre une liste tres complete de references sur ce groupe de plantes. Dans la premiere partie du Plan d’action, des botanistes specialistes des differentes familles, decrivent la repartition, la diversite, les menaces et Mat de conservation des huit principaux groupes taxonomiques de cactus et de plantes succulentes, notamment les Agavaceae, Aizoaceae, Aloaceae, Asclepiadaceae, Cactaceae, Crassulaceae, Euphorbiaceae et Portulacaceae. Certaines de ces familles ont une grande importance economique, principalement dans le cadre du commerce de plantes ornementales, d’autres, telles que les Agavaceae, sont utilisees dans les industries des fibres et des aliments.

Le second chapitre du Plan d’Action decrit et evalue les mesures de conservation des plantes succulentes existant a travers le monde, y compris la reglementation relative a leur commerce et les mesures de conservation in situ ou ex situ. L’objectif de ce chapitre est d’identifier les actions de conservation reussies qui pourraient servir de modeles dans d’autres situations et de mettre en evidence des priori& pour la poursuite des activites de conservation. En analysant les mesures de conservation prises au niveau international, il est particulierement important de montrer comment les exigences de la conservation des plantes succulentes peuvent etre integrees dans le cadre plus general de la conservation de la biodiversite.

Les actions de conservation pour les plantes succulentes doivent etre planifiees en priorite au niveau national et local et etre realisees autant que possible par des acteurs des pays concern& et des experts locaux. Ce principe s’applique a tout le Plan d’action et, par consequent, dans le troisieme chapitre du Plan d’Action, les comptes rendus regionaux, ont ete prepares principalement par des experts des regions concernees. Ce troisieme chapitre met l’accent sur les regions du monde qui sont les plus riches en plantes succulentes.

Le dernier chapitre du Plan d’Action decrit les actions de conservation prioritaires proposees a l’echelle mondiale par les membres du Groupe de specialistes des cactus et des plantes succulentes. Basee sur nos connaissances actuelles, la mise en oeuvre de ces propositions permettra de sauvegarder la plus grand diversite possible de plantes succulentes.

En conclusion, les recommandations du Groupe de specialistes des cactus et des plantes succulentes de la CSE sont les suivantes :

l Effectuer des recherches sur le terrain pour verifier la taxonomie et l’etat de conservation des plants succulentes,

l Ameliorer la protection in situ des plantes succulentes en developpant le reseau d’aires protegees,

l Coordonner la conservation ex situ des especes menacees de plantes succulentes pour renforcer, quand cela est possible, leur conservation dans leur habitat nature&

l Mettre en oeuvre des reglementations nationales efficaces pour toutes les especes de plantes succulentes menacees,

l Controler de man&e stricte le commerce de toutes les plantes succulentes sauvages menacees par le commerce international,

l Sensibiliser le public et les autorites sur la valeur des plantes succulentes, le besoin de les conserver pour le futur et l’importance de leur utilisation durable.

ix

Acronyms used in this Action Plan International Plant Genetic Resources

Institute

International Succulent Institute

International Organization for Succulent Plant Study

International Union for Conservation of Nature and Natural Resources - The World Conservation Union

Jardin Botanic0 National “Rafael M. Moscoso” [Dominican Republic]

Universidad National Autonoma de Mexico

Viceministerio Forestal, Ministerio de Agricultura [Cuba]

Museo National de Historia Natura [Cuba]

National Botanical Institute [South Africa]

Natural History Society [Jamaica]

Natural Resource Conservation Department [Jamaica]

National Trust for the Cayman Islands

The New York Botanical Gardens

Instituto Politecnico National, CIIDIR [Mexico]

Reconstruction and Development Programme [South Africa]

Repertorium Plantarum Succulentamm

Southern African Development Community

Secretaria de Agricultura y Recursos Hidraulicos [Mexico]

Secretaria de Desarrollo Social [Mexico]

Secretaria de Medio Ambient, Recursos Naturales y Peces [Mexico]

Universidad Autonoma de San Luis Potosi, [Mexico]

Protocol on Specially Protected Areas and Wildlife for the Caribbean Region of the Cartagena Convention

Species Survival Commission [ IUCN]

Trade Records Analysis of Flora and Fauna in Commerce

Universidad Autonoma de Tamaulipas [Mexico]

United Nations Commission on Environment and Development

United Nations Educational, Scientific and Cultural Organization

United States Fish and Wildlife Service

World Conservation Monitoring Centre [United Kingdom]

World Wide Fund for Nature

Stadtische Sukkulenten-Sammlung Zurich [Switzerland]

AGUAT

AIAS

ANAE

Universidad de San Carlos [Guatemala] IPGRI

Italian Succulent Plant Society

N Environmental ational Association for Actions [Madagascar]

IS1

10s

Agence Nationale pour la Gestion des Aires Protegees [Madagascar]

ANGAP IUCN

Australian Network for Plant Conservation

Universidad Autonoma de Baja California [Mexico]

ANPC

BCMEX

Botanical Gardens Conservation International (formerly BGCS)

JBN-RMM

BGCI MEXU

CAMPFIRE Corn munal Areas Managem for Indigenous Resources

ent Programme MINAGRI

CBHL The Council on Botanical and Horticultural Libraries MNHN

NBI

NHS

NRCD

Centro de Education Ambiental [Puerto Rico]

Instituto de Historia Natural de Chiapas [Mexico]

Centro de Information Ambiental de1 Caribe [Puerto Rico]

Centro de Investigaciones Biologicas [Mexico]

Centro Interdisciplinario de Investigation para el Desarrollo Integral [Mexico]

Convention on International Trade in Endangered Species of Wild Fauna and Flora

CEA

CHIP

CIAC NTCI

NYBG

OAX CIB

CIIDIR RDP

CITES RPS

SADC

SARH Comision para la Protection de la Flora, la

Fauna y el uso rational de 10s Recursos Naturales [Cuba]

La Comision National de la Biodiversidad [Mexico]

Consejo National de Ciencias y Technologia [Mexico]

COMARNA

SEDESOL

SEMARNAP CONABIO

SLPM CONACYT

SPAW CPC

CPD

DEF

DES

DNP

Center for Plant Conservation [USA]

Centres of Plant Diversity

Direction des Eaux et For&s [Madagascar]

Desert Botanical Garden [USA]

Direction National de Parques [Dominican Republic]

Departamento de Recursos Naturales [Puerto Rico]

ssc TRAFFIC

UAT DRN

UNCED EC

FSA

HNT

IABG

European Community

Flora of southern Africa region

Huntington Botanic Gardens [USA]

International Association of Botanic

UNESCO

Gardens USFWS

WCMC

IBUG

IES

INE

Universidad de Guadalajara [Mexico]

Instituto de Ecologia y Sistematica [Cuba]

Instituto National de Ecologia [Mexico]

WWF

zss

Introduction

Succulent plants have a global distribution and are of Conduct has been widely publicised through nationalrepresented in nearly all habitat types. Over 30 botanical cactus and succulent societies and has provided a modelfamilies have succulent plant species, ranging from tiny for use by other specialist plant societies. The Code ofannual plants to huge trees. The Cactaceae is the largest Conduct was updated in 1990 (Oldfield 1990).and perhaps the best known of the succulent plant The 10s Conservation Section provided the basis forfamilies. The unifying characteristic of succulent plants is the formation of the IUCNSSC Cactus and Succulenttheir ability to store water in one or more organs of the Specialist Group in 1984, and the two groups continue toplant, giving rise to stem succulents, leaf succulents, and work together sharing a common membership and regularroot succulents - the so-called caudiciform plants. meetings. The SSC Cactus and Succulent Specialist

“A succulent (or succophyte) is a plant possessing at least one succulent tissue. A succulent tissue is a livingtissue that, besides possible other tasks, serves and guarantees an at least temporary storage of utilisablewater, which makes the plant temporarily independent from external water supply when soil water conditionshave deteriorated such that the root is no longer able to provide the necessary water from soil.” von Willert et al.(1992)

Succulents, with their often bizarre growth-forms andattractive flowers have long attracted the attention ofbotanists, both amateur and professional, andhorticultural enthusiasts. Despite the widespread interestin succulent plants, and their ecological and economic

Strombocactus disciformis, Mexico.

importance, relatively little attention has been paid totheir conservation needs, outside the confines of thespecialist societies. The International Organization forSucculent Plant Study (10s) has been instrumental inpromoting conservation mainly through the publication ofa Code of Conduct. First published in 1973, the 10s Code

Group has undertaken important conservation fieldworkin Chile, Mexico and Ecuador, has acted as an expandednetwork for conservation discussion and exchange ofinformation, and has contributed to CITES matters. Theidea for an SSC Action Plan for cacti and succulents wasfirst discussed by the Specialist Group in 1990 at theGroup’s meeting held in Bonn, Germany. There wasgeneral agreement that preparation of the Plan shouldproceed as rapidly as possible. There is a sense of urgencyin all plant conservation matters and particularly so forgroups of plants where information, although by nomeans complete, is sufficient to prioritise and initiate longoverdue conservation activities.

Arrangements for the preparation of the Action Planwere confirmed in 1993 at the Group’s meeting in Maltaand a first draft was prepared in time for the 1994 Groupmeeting, coincident with the 23rd 10s Congress and 24thAETFAT Congress held in Wageningen. The taxonomicaccounts for the Action Plan were prepared in 1993together with the general sections on legislation andcontrolling the trade. Preparation of the regionalaccounts, which generally necessitated informationgathering from diverse sources and wider consultationamongst local organisations and experts, was finalised atthe end of 1995. The Wageningen meeting provided aparticularly useful opportunity to discuss the African andMadagascan accounts for the Action Plan, and the CITESPlants Committee meeting held in San Miguel deAllende, Mexico in May 1994, provided an opportunity toreview the Mexican account. The timespan involved inpreparation of the Action Plan already means thatupdates will be necessary for certain sections with regardparticularly to taxonomic references now published andCITES trade information. However, the Group’s mainpriority, following this period of information gathering, isnow to implement the Action Plan.

1

The Action Plan aims to bring together currentinformation on the conservation of cacti and succulentsfrom around the world in order to provide a summary ofthe present situation. This provides the backgroundagainst which to set priorities for conservation action. Inpreparing an overview Action Plan of this nature it hasnot proved possible to be fully comprehensive in thespecies and areas covered. The north temperate regionsof Europe and Asia, for example, which are not coveredspecifically in the Action Plan, are by no means withoutsucculents but do not have the same diversity, degree ofendemism, and urgency for succulent plant conservationas, for example, Madagascar, Mexico, and Namibia. TheSpecialist Group is keen to learn of other succulent plantspecies and other areas of succulent plant diversity whichare in need of conservation attention and also of ongoingconservation initiatives to which it can contribute.

The annexes to this Plan provide various regionaltaxonomic lists of succulents. Unless otherwise noted, theconservation status given for each taxon follows the IUCNRed List categories. With the exception of the BrazilCactaceae reported by Nigel Taylor, all of the lists followthe old criteria (pre-1994). Work is ongoing by specialiststo apply the new criteria which offer a more objective anddetailed evaluation of these species’ threat of extinction.Annex 16 details both versions of the IUCN Red Listcategories.

A precise definition of the term ‘succulent’ has notbeen attempted for the purposes of the Action Plan andsome doubtfully succulent species have been includedwhere these are of conservation concern. Some of the

caudiciform plants, for example, are marginally succulent,but are fashionable at present with succulent plantcollectors and subject to trade pressures. Cycads andsome orchids can also be considered succulent, but theseare not included in the Action Plan because they fallwithin separate SSC Action Plans (for Orchids seeIUCN/SSC Orchid Specialist Group 1996; IUCN/SSCCycad Action Plan, in prep). Certain genera of theBromeliaceae are included within the Action Planbecause plants of this family often have similar lifeforms,grow in similar habitats, and face the same threats asplants more commonly treated as succulents. It is,however, hoped that the Bromeliaceae will soon have anSSC Conservation Action Plan of their own.

Further field research and taxonomic studies areurgently needed as a preliminary conservation activity forsucculent plant species is various parts of the world. Theneed for further information should not, however, hold upthe implementation of in situ and ex situ conservationactivities as outlined in the Action Plan. Members of theSSC Group have expertise in succulent plant taxonomy,ecology, conservation planning, conservation legislation,information management, development of recovery plansand cultivation of succulent plants. Integration ofdifferent approaches and methodologies for conservationwill be important to ensure the conservation of maximumsucculent plant diversity. The preparation of the ActionPlan is not an end in itself but a beginning. The SSCCactus and Succulent Specialist Group is committed toimplementation of the Plan and offers its assistance to allwho are in a position to take the action proposals forward.

Chapter 1

Taxonomic Groups

AgavaceaeWendy Hodgson

The Agave family (Agavaceae Endlicher) is a group ofeconomically important succulent plants with a naturaldistribution in the drier regions of the tropics andsubtropics. The botanical limits of the family areundecided, but for the purposes of this Action Plan theAgavaceae is considered to comprise 18 genera andapproximately 625 species. Many of these species remainpoorly known in the wild. Herbaria and field surveys areurgently needed to determine conservation status for thespecies. Despite lack of detailed field information,progress has been made towards developing aconservation programme, particularly for the Americanspecies, for example through the SSC Agavaceae ActionPlan Workshop held at the Desert Botanical Garden,Phoenix, in 1992.

agaves in this respect while other genera now consideredin Agavaceae (Nolina and Dracaena) do not. Cronquist(1981) considered the Aloaceae and Agavaceac asparallel derivatives from the Liliaceae with only slightdifferences.

McVaugh (1989) points out that unless some basisother than plant habit can be found, it is impossible todistinguish between the Agavaceae and Liliaceae in theNueva Galicia flora of western Mexico. In Manfreda,Polianthes, and Prochynanthes, genera generallyconsidered to be closely related to Agave, plants areherbaceous, commonly have fleshy roots developed froma short rhizome, and have soft, thin, slightly succulentleaves which in most species die back annually; marginalteeth, if present, are soft. McVaugh (1989) suggests onesolution might be to include all herbaceous genera in theLiliaceae, and only the more woody groups in Agavaceae.However, the problem is more complicated in that there

Box 1 .I Botanical characteristics of AgavaceaeThe Agave Family is characterised by stout, simple or sparingly branched, arborescent shrubs (or sometimes trees), orshort-stemmed, somewhat herbaceous plants with a short rhizome or erect caudex; leaves simple, alternate, sessile,tending to be crowded in dense rosettes at ends of stems or branches or at ground-level on a short stem, generallythickened, leathery or firm-succulent (in contrast to soft-succulent as in Aloaceae), often prickly on margins and spine-tipped; flowers in dense racemes or panicles or heads terminating the stem (plants are monocarpic, as in Agave), oraxillary and subterminal (plants are polycarpic, as in Yucca); perfect, sometimes unisexual, perianth consisting of tepalsarranged in 2 whorls of 3, petaloid, often thick and fleshy, distinct or fused below to form a tube, stamens 6, filamentsdistinct, fused to tepals or base of tube, ovary superior or inferior, 3-carpeled, usually with nectaries style usually terminalwith 3 stigmas, ovules l-many, fruit a loculicidal capsule or berry, seeds flattened; chromosome counts vary from x= 16-30+.

Systematic treatment

The Agavaceae has undergone many changes since it wasproposed by Endlicher in 1841. Cronquist (1981)maintains the Agavaceae as a family distinct from theLiliaceae and Amaryllidaceae based on the specialisedgrowth habit. Agavaceae are characterised by being stout,simple or branched shrubs or trees, or herbaceous plantsarising from a caudex, often forming succulent rosettes asopposed to Liliaceae and Amaryllidaceae which areherbaceous perennials, usually dying back to the ground.Yucca and Agave share the karyotype of 20 small and 5large chromosomes. This trait was considered unique atone time and provided the incentive to remove Yuccafrom the Amaryllidaceae to the Agavaceae. However, thepresence of few large and many small chromosomes ismore common throughout the Liliaceae than oncethought. Hosta, a genus with a very different habit andincluded within the Liliaceae, resembles yuccas and

are obvious similarities (and presumed relationships)between Agave and Manfreda. In addition, Manfreda isclosely related to Polianthes and Prochynanthes. He thenjustifiably asks that if Agave and Manjreda, and Manjredaand Polianthes, are so closely related that they could evenbe considered congeneric, then can Agave and Polianthesbe assigned to different families?

Various subdivisions of the family have been made.Hutchinson (1934) divided Agavaceae into six tribes:Yucceae (Yucca and HesperaZoe), Dracaeneae (Cordyline,Cohnia, Dracaena, and Sansevieria), Phormieae(Phormium), Nolineae (Nolina, Calibanus, Beaucarnea,and Dasylirion), and Polyantheae (Polianthes,Prochynanthes, Bravoa, and Manfieda). Cronquist (198 1)also recognised 18 genera but conceded that so treatedthe Agavaceae may not form a natural group. Phorrniumand Doryanthes, usually included in this family, may notproperly belong with other genera. Serologically Agaveand Yucca seem to stand apart from a group consisting of

Dracaena, Nolina, Sansevieria, and Cordyline. Dahlgren etal. (1985) recognise Agavaceae with eight generasubdivided into two subfamilies: Yuccoideae (Yucca andHesperaloe) and Agavoideae (Agave, Manjreda, Polianthes,Prochynanthes, Beschorneria, and Furcraea), based oncytological, anatomical, and embryological studies.

Recent studies at the University of Texas, Austin,support recognising the Nolinaceae as a monophyleticgroup including, among others, Beaucarnea, Nolina,Dasylirion, and Calibanus (see Hernandez and Simpson1992). Recent s tudies on the molecu la r andmorphological phylogeny of the Agavaceae by DavidBogler at the University of Texas support the treatment ofHutchinson and Dahlgren and the findings of Hernandezand Simpson. Preliminary work, which includedchloroplast DNA restriction site analysis, suggests thattwo distinct major lineages occur (D. Bogler, pers. comm.,1994). The first group includes Hosta, Camassia, andAgavaceae. Within the Agavaceae are the seeminglyclosely related Agave, Manfreda, Polianthes, a n dProchynanthes; Beschorneria and Furcraea appear moreclosely related to the more primitive Agave dasylirioidesand A. striata, while Yucca appears to be more closelyrelated to Camassia than to Agave. The second lineageincludes the Convallariaceae, Dracaenaceae andNolinaceae, the latter a monophyletic group includingNolina, Dasylirion, Calihanus, and Beaucarnea. In thisgroup, Calibanus appears most closely related toBeaucarnea.

In this account, the Agavaceae is recognised sensuZato, following Cronquist’s more inclusive, albeitunnatural treatment. Most scientists, at least in the USA,recognise this treatment and the Flora of North Americaincludes Dasylirion in the Agavaceae. As defined here, theAgavaceae includes 18 genera and approximately 625species.

Distribution and diversity

Naturally distributed through the American andCaribbean arid and tropical climes, agaves and theirrelatives have been widely dispersed since 1492 to othercontinents, where they are viewed as having considerableeconomic and horticultural importance. The origins ofthese genera and species appear to be related to theevolution of the Neotropical flora in the cordilleras ofMexico and Central America. Best known is theconsiderable adaptive radiation of the genus Agave thathas occurred in the Sierra Madre Occidental of Mexico(Gentry 1972, 1982) undoubtedly associated with thedevelopment of the Madro-Tertiary flora (Axelrod 1958).

Through secondary adaptive radiation, members ofthe Agave family have adapted to habitats ranging fromthe temperate woodlands and prairies of eastern NorthAmerica to the wet tropics and subalpine habitats ofSouth America. Nevertheless, most species in the familyoccur presently in arid, semi-arid, and dry subtropicalvegetation zones in both coastal plains and adjacentmontane landforms. The predominance of CAMmetabolism and succulence among these species allowsthem to remain dormant during extended seasonaldroughts and periods of high temperatures. Theextremely high water use efficiencies that have beenrecorded among agave cultivars (Nobel 1988) are alsocharacteristic of the wild species and not simply a productof domestication.

Of the 18 genera in the Agavaceae, species richness ishighest in the genus Agave (200+ species) and thendescends in roughly the following order: Pleomele (140,Old World tropics and subtropics, if treated as distinctfrom Dracaena), Dracaena (50, Old World tropics),Sansevieria (50-60, South Africa, Madagascar, andArabia), Yucca (35-40, south USA, Mexico, Guatemala,and Cuba), Nolina (25-30, south-west USA to central

Agave collection, DesertBotanical Garden,Arizona.

Collection of centre of Agave plant for fibres, Pehita,Tamaulipas, Mexico.

Mexico), Manfreda (25, south and south-east USA toHonduras), Furcraea (c. 20, central Mexico to SouthAmerica), Dasylirion (14-17, south-west USA to Oaxaca),Cordyline (10-15, Old World tropics), Polianthes(including Bravoa 20, Mexico), Beschorneria (10, Mexicoand north Guatemala), Beaucarnea (9, Mexico to SouthAmerica), Hesperaloe (4, central Texas and northMexico), Doryanthes (3, Australia), Phormium (2, NewZealand), Prochynanthes (1, Mexico), and Calibanus (1,Mexico).

The following centres of Agavaceae diversity havebeen identified by Gentry (1982) Reichenbacher (1985)and Garcia-Mendoza (1987, 1989): the Apacheanwoodlands and grasslands of the Deming Bridge betweenthe Sonoran and Chihuahuan Deserts; the subtropicalCape region of Baja California; the northern and centralSierra Madre Occidental, including temperate and semi-arid woodlands and dry subtropical thornscrub; theecotone between the semi-arid highlands of centralMexico and the wetter Sierra Madre Oriental; the moretropical montane belt of ranges in Michoacan andadjacent states; the Chiapan and Oaxacan highlands ofsouth-western Mexico and adjacent Guatemala, includingsemi-arid valleys in the rain-shadow of the Sierras, andadjacent humid uplands. Oaxaca has the greatest diversityof Agavaceae in the Mexican Republic with 42 taxa.

The species list of Agavaceae in Annex 1 indicatesspecies which are currently considered to be narrowendemics, inhabiting zones of less than 100 km in lengthand width (10,000 km2). Here high levels of endemism inareas of high species diversity are expected. Indeed, of the42 taxa in the State of Oaxaca, Mexico, 32 are endemic tothe Republic of Mexico and six are endemic to the state.Hybridisation, polyploidy, and vegetative reproductionare important processes in species formation in Agave(Pinkava and Baker 1985), and probably other membersof the family. For example, Yucca campestris, a sand-duneendemic of south-western Texas, may have originated

from hybridisation between Y. elata and Y. constricta(Powell 1988). Peninsular floras such as that of BajaCalifornia also contain high levels of endemism. Of the 25taxa of Agave that occur in Baja California, 20 areendemic (Gentry 1972).

Species richness is also affected by the local diversityand abundance of their pollinators; for example, Oaxacahas nine species of nectar-feeding bats specialising onAgave (A. Garcia-Mendoza and H. Arita, pers. comm.).The interactions between certain members of theAgavaceae, e.g. Agave, Manfreda, and Yucca, and theirpollinators Leptonycteris and Tegeticula, have been widelydiscussed as classic examples of pollination syndromesbetween mutualistic species which generate and maintaincertain aspects of the structure of their bioticcommunities (Webber 1953; Howell 1974; Arita andMartinez de1 Rio 1990). Several genera of nectar-feedingbats, bees, hummingbirds, wasps, moths, and hawkmothsmay have co-evolved mutualistic relationships with someagaves; yucca moth co-evolution with Yucca species hasalso been documented in intricate detail. The populationreduction or local extirpation of these pollinators maylead to reduced fertility and seed set in individual plants(Equiarte and Burquez 1988) but there is as yet noevidence that populations of long-lived polycarpicperennials have been severely reduced (Nabhan andFleming 1992) . However , r e in t roduc t ion andtranslocation efforts of species in the Agavaceae are likelyto fail if pollinators are no longer present in sufficientabundance to sustain populations.

Local uses and commercialisation

The Agavaceae is of considerable economic importance.Fibrous leaves provide cordage and are used in makingmats, baskets, hats, thatches, paper, fans, sacks (Nohna,Dasylirion, Beaucarnea, Agave, Hesperaloe f’unifi’ra,Phormium, Furcraea, Sansevieria, Yucca), flowerarrangements (Agave), brooms (Dasylirion), and packsaddles (Furcraea). Trunks, stems, and flower stalks

Cut Agave lechuguilla, San Luis Potisi, Mexico.

provide food for humans and cattle (Agave, Dasylirion,Nolina) and alcoholic beverages (Agave, Dasylirionincluding D. cedrosanum, Yucca). Flowers of Yuccaelephantipes are eaten by people in south-east Mexico.Fruits provide food for humans (the baccate-fruitedspecies of Yucca, including the more restricted Y.endlichiana, Y arizonica, Y. grandiflora, and Y. madrensis).Trunks provide fuel (Dasylirion), while trunks and oldstalks are used for the construction of temporary shelters,posts, porches, and corrals (Agave, Dasylirion, Nolina,Yucca). Some taxa are grown as living fence rows (Agaveand Yucca). Rhizomes (usually erroneously referred to asroots) and, to a lesser extent, leaves, were or still are usedto produce soaps, shampoos and detergents (Yuccaincluding Y. madrensis, Manfreda, Prochynanthes,Polianthes, Furcraea longaeva, and Nolina palmer-i, a BajaCalifornia endemic), as insect repellents (Prochynanthes),a n d a s medicines (Ag a ve, Yucca, Manfreda,Prochynanthes) . Flowers are used in perfumery(Polianthes tuberosa, known only from cultivation).Flowers of Polianthes tuberosa are used as cut and gardenflowers while other species of Pohanthes are sometimessold as cut flowers in Mexican markets. Many taxa arenow grown as ornamentals: Agave, Calibanus, Dasylirion,Doryanthes, arborescent Nolina, Hesperaloe, Yuccaincluding the Joshua tree Y. brevifolia and Y. rostratacollected from the wild for landscaping, Manfreda,Pleomele, Polianthes, and Beaucarnea.

Rare and threatened taxa

It is difficult to determine whether specific taxa within theAgavaceae are threatened, or if so, to what degree.Despite recent field work, taxa are still poorly known orcollected, particularly in south-central Mexico. Inaddition, interspecific hybridisation is a relativelycommon phenomenon within the family and there occursmuch variability within and intergradation between

Machine preparing Agave fibres, Ecuador.

populations. As a result, defining taxa becomes moreproblematic and adds to the difficulty in determiningrarity.

In the Republic of Mexico the native habitat for sevenpercent of Agave species is unknown, while 15 percent areknown from three or fewer herbarium collections at thetime of Gentry’s (1982) monograph (Reichenbacher1985). Approximately 50 percent of the entire genus iseither poorly known, potentially rare, or occur as fairlyrestr icted endemics (Annex l), and therefore ofconsiderable conservation concern. Of the 25 knownspecies of Manfreda, 14 have limited ranges or fewcollections. Of these 14 species, six are known from six orfewer collections. Of the 13 species of Pohanthes at leastnine have limited ranges within which populations arescattered (S. Verhoek, pers. comm. 1993). Until there is arevision of Furcraea identification of all species (with theexception of F. bedinghausii) is doubtful. Authors point

Making cordage ofAgave fibres, Ecuador.

out that some Furcraea species have limited ranges, withsome native to only a few Caribbean Islands. At the timeof Standley’s (1920) text, of the 22 or so species of Nolina,five were known only from their type localities and sevenwere endemic to a state or small part of a state. Likewise,of the approximately 17 species of Dasylirion, five wereknown only from their type localities and an additional sixwere endemic to one state. Subsequent studies ofDasylirion will result in taxonomic changes and rangeextensions for Mexican species; in addition, it appearsthat none of the (approximately 10) well defined speciesare rare (D. Bogler, pers comm. 1993), although they maystill be endemic to one state. All members of the genusHesperaloe are considered to have limited ranges, with theexception of H. funifera. Hernandez (1993) considers thewhole genus Beaucarnea as threatened, with five speciesbeing particularly sensitive.

Regionally, Agavaceae taxa are often poorly known orlimited in distribution. For example, over 30 percent ofthe agaves in Oaxaca are represented by only a fewspecimens. Little or no information is available for sevenAgave species endemic to the Bahamas and they may bevery rare. Within Guatemala, eight species of Agave, onespecies of Beaucarnea, and three species of Furcraea areconsidered to have limited distributions.

Recognised threats

Threats to agave family members can be classified asglobal (climate change, air contamination), habitat-specific (overgrazing, land conversion, competition withexotics), taxon-specific (vulnerability to introduced pestsand diseases, loss o f po l l ina tors , economicoverexploitation), and population-specific (collection byhobbyists, depletion for local, traditional uses). Becausemany species in the agave family are characterised eitherby few populations, low numbers of individuals perpopulation, or both of these factors, they are vulnerableto environmental and demographic stochasticities. Inaddition, little information is available on the majority ofspecies within the family thereby reducing the chances ofappropriate management of their populations.

By far, the three most common pressures onAgavaceae taxa are: 1) land clearance and conversion ofnative vegetation for agriculture; 2) direct and indirecteffects of overgrazing by livestock; and 3) overcollectionfor ornamental purposes, and to a lesser extent, fibre andalcohol. For example, the very beautiful Agave wercklei isendemic to the Rio Grande region on the Pacific slope ofCosta Rica. At present, their populations are beingadversely impacted by the development of coffeeplantations (A. Mendoza, pers. comm. 1993).

Livestock, particularly goats, eat young plants andemerging flowering stalks which limits sexualreproduction and reduces genetic variation. Agavepeacockii is known from only a small area in the vicinity ofTehuacan, Puebla, where Gentry (1982) observed only 40-50 plants in the mid-1960s. Local people believe it has the

Fortunately, with the availability of commercialalcoholic beverages, the desire and need to produce‘bootleg’ alcohol from Agave and Dasylirion have becomeless important. There have, however, been dramatic localdeclines in Agave abundance in parts of Sonora whereAgave angustifolia, A. palmeri, and A. shrevei have beenintensively harvested for illicit mescal production (G. P.Nabhan, pers. comm. 1993).

Overcollection for ornamental purposes continues topose serious threats to various genera within theAgavaceae. The fact that all species of Beaucarnea areconsidered threatened, mainly by overcollection, raisesserious concerns. Illegal collectors have made it profitablefor campesinos to collect material which is said to go toeither the United States or Germany (Hernandez 1993).Hernandez (1993) points out that overcollection of seedsand seedlings are affecting natural repopulation of thespecies. As members of the genus are dioecious,overcollection of young and adult plants is changing thesex ratios and reducing the production of seeds, therebylowering the size and genetic variability of thepopulations. The need to regulate the export of these andother overcollected plants is critical. The interest anddemand for particular genera, including Aglve,Hesperaloe, Yucca, and Beaucarnea, in the horticulturaltrade has increased dramatically over the last 10 years andis expected to continue. However, only five out of over180 potentially rare or restricted taxa within theAgavaceae sensu Zato are listed in the Appendices ofCITES.

Actions for conserving genetic diversity

There are four current activities which directly orindirectly conserve the genetic diversity found within thisfamily: 1) efforts to resolve the causes of threats tohabitat; 2) efforts to protect selected habitats where rareor otherwise threatened taxa occur through establishingbiosphere reserves, sanctuaries, parks, or other protectedareas where their populations can be maintained,translocated, or enhanced; 3) efforts to establish andenforce laws limiting the extraction from the wild andtransport and commerce of rare, threatened, orvulnerable taxa; and 4) efforts to establish ex situ samplesof the genetic variation of each species for propagation inbotanical gardens, in vitro laboratories, or for storage asseeds or tissue cultured germplasm in gene banks. Onlythe Desert Botanical Garden, Phoenix, has developed agene bank specialising in the Agavaceae, although over 40botanical gardens contain CITES-listed endangeredspecies from this family. The principal ex situ collectionsof the agave family occur at the Desert Botanical Garden

Box 1.2 Ex situ conservation of Agavaceae at Desert Botanical Garden, PhoenixDuring 1991 and 1992, a systematic review of the Agavaceae collection was carried out at the Desert Botanical Gardenas part of an Institute of Museum Services grant (Ecker and Burgess 1992). The review and subsequent report addressedI) the determination of the scientific value of the Agavaceae collection and verification of the identity of as many speciesas possible, 2) identification of rare species in the collection which were noted in the database, 3) creation of a priority listof species to be propagated, and 4) development of a Long Range Conservation Plan which identified species requiringpropagation and/or acquisition. Strategies for propagation and collection of plants were developed on a species-by-species basis. A conservation plan outlining the status of each taxon in the Agavaceae collection and a plan for the longterm conservation of each taxa were developed. Genetically representative seed collections have been established fromseveral populations of Arizona taxa. Pollen is collected and stored from live plant material for controlled pollination. Seedproduced from controlled cross-pollination is made available to other institutions via the Index Seminum program.Although still improving, the Desert Botanical Gardens ex situ program for Agavaceae can provide a template for otherinstitutions considering the development of such a program.

Arizona), the Jardin Botanic0 of the UniversidadAutonoma de Mexico (UNAM) (Mexico City), andL

huntmgton Botanical Garaen (Camorma).

AizoaceaeSteve HammerT 1. 1 - 1 . lfl 1 /r( 1.r * \

With regard to significant in situ conservation effortswhich include rare or threatened populations ofAgavaceae, the following UNESCO-recognised biospherereserves contain several species: Ranch0 El Cielo,Tamaulipas; Bolson de Mapimi, Coahuila; Big Bend,Texas; El Pinacate, Sonora; Isla Tiburon, Sonora;Desierto de Vizcaino, Baja California; La Michilia,Durango; Sierra de Manantlan, Jalisco; and SelvaLacandona, Chiapas. None of these protected areas, asfar as is known, has developed action plans specific tomanaging and conserving local members of theAgavaceae.

A conservation strategy for the Agavaceae of centralMexico to northern South America was presented at theSSC Agavaceae Action Plan Workshop held at the DesertBotanical Garden, Phoenix in 1992 (Mendoza and Bye1992). The strategy is incorporated into the ActionProposals in Chapter 4 of this Action Plan.

The author would like to acknowledge the following people whocontributed to the preparation of this account for the Action Plan:Dr Edward F. Anderson, Alfonso V. Banuet, Miquel ChazaroBasenez, David Bogler, Dr Tony Burgess, Joe Clements, LizSlausen (Ecker), Dr Richard Felger, Gary Lyons, Dr BruceMacBryde, Enrique Martinez, Nora Martinez, Abisai GarciaMendoza, Esther S. Monarque, Dr Gary Nabhan, Peggy Olwell,Sue Rutman, Dr Luis Hernandez Sandoval, Dr Susan Verhoek.

The Aizoaceae Rudolfi, the Mesembryanthema, is thesecond largest family of succulent plants after Cactaceae.The family is currently divided into five subfamilies, ofwhich two, Mesembryanthemoideae and Ruschioideae,comprise the bulk of the succulent species within thefamily as a whole. The Mesembryanthemoideae andRuschioideae form a clade known informally asMesembryanthema sensu H.E.K. Hartmann. Only theplants of this group, known colloquially as ‘mesembs’, areconsidered here. The remaining three subfamilies havefew, if any, true succulents and have thus escaped thecollector’s zeal, although they do include plants ofagricultural importance.

For two centuries Mesembryanthema or ‘mesembs’have been intensely studied both by professionals andamateurs, resulting in ‘taxonomic chaos’. Now, throughwork at the University of Hamburg, some accord is beingreached. As presently understood the clade (the formerMesembryanthemaceae Fenzl) comprises some 120genera informally arranged into 15 groups (Hartmann1991) based on fruits, flower, and vegetative characters.

Among the succulent subfamilies the two basicdivisions, Mesembryanthemoideae and Ruschioideae, areseparated by nectary form, by the axile or basal-parietalplacentation of their gynoecia, and by different types ofexpanding structures in their fruits. Mesembryan-themoideae comprises perhaps 100 species in about 10genera. Only a few of its species are suited to or areattractive under cultivation. In the wild, thev do havesome importance for grazing.

Box 1.3 Botanical characteristics of MesembryanthemaMesembryanthema are characterised by a high degree of leaf succulence, and especially by their hydrochastic capsules(more or less persistent dried fruits, remarkable in structure and highly responsive to moisture). Other characteristics arethe (frequent) possession of bladder idioblasts, bright petals (petaloid staminodes) of androecial origin meronectaria,well-developed valve wings in the fruits and a basic chromosome number of x=9. The monophyllum comprises mainlyperennials, some biennials, and a few annuals. In their growth form, species range from tiny geophytes which produceonly small tufts of barely visible leaves annually, to long-lived shrubs 3m tall.

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Ruschioideae, by far the larger group, comprisesabout 110 genera with perhaps 1200 species. Revisions ofthe larger genera are not complete. Many more specieshave been described, especially in the 1920s and 1930s;tautologies resulted from the independence andconflicting concepts of distant authors. As a general rule,the described species will probably be reduced to one-third of their former number. On the other hand, many ofthe less attractive species in both the subfamilies areundercollected, overlooked, and undescribed even thoughmany of them are biologically fascinating. It is probablethat many geophytes and very narrowly endemic dwarfspecies are still unknown. Only a few extinctions,however, have been suspected.

The most compact species in the Ruschioidaeae,members of the so-called highly-specialised or ‘stemless’genera, have attained a degree of diversity and adaptation(such as dimorphic and or fused leaves, fenestration,riddling patterns, hair coverings) unknown in their long-stemmed leafier allies. It is these bizarre forms whichhave attracted so much attention from collectors. Only afew species in Mesembryanthemoideae, chiefly bothspecies of Dactylopsis have attracted as much notice,though some geophytic ‘mesemboids’ might do so oncetheir ornamental potential is better understood.

Centres of diversity

The main centres of diversity occur within South Africaand, to a lesser extent, Namibia. The occurrence ofMesembryanthema in northern Africa and theMediterranean rim involves a few species of naturalorigin; the majority have, however, been introduced intothis area.

Cheiricfopsis derenbergiana, Karoo BotanicalGarden, South Africa.

Dactylopsis digitata, an unusual member of theAizoaceae; its finger-like appearance gives rise to itslocal name of ‘bobbejaans vingers’ or ‘baboonfingers’.

Within South Africa, the highest concentration ofgenera occurs in the so-called ‘Vanrhynsdorp centre’ asdefined by Nordenstam (1969) with 31 genera within onequarter-degree square. The ‘Gariep centre’ has 30 genera,and the ‘Little Karoo centre’ has 27 (Hartmann 1991).The first two centres constitute southern and northernextremes of the area long known as Ncentres are extremely rich in ende

ama aualand.mic species,

All theand it

should be emphasised that many endemics occupy verysmall habitats. This is particularly true for the Gariepcentre, complex and rough in its geology. Many speciesare restricted to single hilltops or hill groups and this, ofcourse, has strong conservation implications.

In many areas mesembs form theis particularly the case in the so-call

dominant flora; thised Knersv lakte (the

Vanrhynsdorp centre) and in some similar areascharacterised by eroded, quartz-covered shales. In suchareas the soils are poor in humic content and often high insalts. Many habitats are extremely sensitive andspecialised. Unfortunately, many of the more peculiarhabitats also harbour valuable mineral deposits such ascopper, nickel, titanium, diamonds and gold, and mininghas led to the eradication of several plant formations.

Trade

Within the Mesembryanthema, the ornamental speciesare the most important economically. Creepy or shrubbymesembs excel as ground cover plants in temperate areas;some, such as DeZospenna spp., are exceptionally hardy,and some species are wonderfully colourful beddingplants. The highly succulent species in Lit/lops andConophytum are valued as pot plants, though world salesare small compared with figures for Cactaceae.

Illicit trade in field-collected plants is hard to measure,but it seems that the current market is small. This mightbe partly due to a virtuous green consciousness among

9

sellers and collectors, but it can also be attributed to thefact that mesembs are not currently popular orfashionable with succulent collectors. The strongest factoris that almost all ornamental mesembs are now cheaplyavailable as seeds, seedlings, or cuttings. The price ofConophytum burgeri, a peculiar endemic, fell fromUS$300 per wild-collected plant in 1978 to US$3.50 for afive-year old nursery seedling in 1992. It is the intention ofcertain specialist nurseries to widely propagate them, ineffect driving prices down, thus eliminating most of themarket for wild-collected plants.

Threats

The mountainous areas of Namaqualand are notamenable to cultivation, but they are grazed by goatswhich have a damaging impact on the succulent plantflora. The fertile and comparatively moist southern Cape,home to some of the more tender mesembs, including thefynbos species, has experienced great destruction fromagricultural activities. This has had the greatest effect onAloaceae, but mesembs have not entirely escaped, whichmight account for the apparent extinction of Circandra.The greatest threats to the mesembs are certainly thosewhich are hardest to deflect: farming, mining, andurbanisation.

The threat of collecting is generally adequatelycontrolled. In many respects the most attractive plants arefortunate: all can be easily and quickly propagated fromseed, and almost all are already in cultivation in the USAand the UK. Moreover, most plants grow rapidly, so theappeal of the big, old wild plant is absent. The activities ofthe few private field collectors are, however, notoriouslydifficult to control. The nature conservation laws in SouthAfrica are very stringent, but easily circumvented byanyone with the requisite boldness, as the country is toolarge for adequate policing. The best hope is to convincecollectors that “digging them up” damages the plants thatthey claim to love. If specialists are to glamorise andpopularise the plants by publishing attractive articles andtempting descriptions, they must simultaneously provide alegal quick and convenient method for distributing seedsand seedlings from material already in horticulture.

Protection

CITES protection is probably not appropriate for rarespecies of mesembs such as Muiria hortenseae. Thisparticular species is threatened in habitat, occupying atiny but conspicuous roadside site of about one hectare.Trade controls on Muiria could arguably increase thenotoriety and market value of a species which is alreadyovervalued. Furthermore, a well-grown Muiria seedlingwould be impossible to distinguish from a wild plant.Purchase and preservation of the most sensitive tracts ofland may be the only option to protect mesembs in theKnersvlakte region.

AloaceaeErnst J. Van Jaarsveld and Gideon F. Smith

Although distributed through a large part of the OldWorld, the Aloe family, a fairly small taxonomic group ofmonocots, has its major centre both in evolutionary depthand in total number of species and genera in southernAfrica. Of the approximately 436 species in about sevengenera in the family, there are some 233 species in sixgenera in the area covered by the Flora of southern Africa(FSA) project. This project aims to provide a taxonomicaccount of all known indigenous and naturalised planttaxa of the subcontinent south of, but excluding, Angola,Zambia, Zimbabwe, and Mozambique. In terms ofnumber of species, the Aloaceae is more or lesseighteenth largest family in the subcontinent, and thesixth largest monocot family (cf. Table 4 in Goldblatt1978, and assuming fragmentation of the Liliaceae intoseveral smaller families). However, on a world scale theAloaceae is the largest monocotyledonous family ofsucculent plants and the second largest succulent plantfamily in southern Africa (cf. Asphodelaceae in Tables 1

10

Aloe helenae, listed on CITES Appendix I.

and 2 in Smith et al. 1993, excluding Bulbine andBulbinella, but including Lomatophyllum). The family hasboth horticultural and medicinal value. Pharmaceuticalproducts derived from especially Aloe vera and A. feroxare widely used (Kent 1980; Bloomfield 1985).


From a modest beginning in 1753 as a single lillioid genus,Aloe (1753), this taxon rapidly gained more genera andspecies. In recent years the Liliaceae, where the alooidgenera have been traditionally classified, has beensubjected to significant taxonomic reassessment in termsof its constituent infrafamilial taxa. The differentinterpretations of the circumscription of the Liliaceae byvarious taxonomists have resulted in, amongst others, thetribe Aloineae sensu Hutchinson (1959) being removedfrom the family (Dahlgren and Clifford 1982; Dahlgren etal. 1985). However, the circumscription of the Aloineae,one of 28 tribes recognised by Hutchinson (1959) in theLiliaceae, has undergone comparatively little change. Themajor controversies surrounding the taxonomy of theAloineae have rather centred on genus and speciesconcepts. In line with the classificatory interpretation ofBrummitt (1992) of this natural group, we regard thealooid genera as warranting segregate familial status inthe Aloaceae. The family includes the following sevengenera: Aloe (333 species; Reynolds 1966, 1982),Haworthia (68 species; Bayer 1982) Gasteria (16 species;Van Jaarsveld 1991), Lomatophyllum (12 species;Jacobsen 1986) Astroloba (7 species; Roberts Reinecke

1965), and two monotypic genera, Chortolirion (Smith1991b) and Poellnitzia (Smith and Van Wyk 1992a). Thegeneric status of the two monotypes has been disputedfrom time to time and they are sometimes included inHaworthia. Although the boundaries of many of thealooid genera and some of the species are still beingdebated, general agreement exists over the question offamilial monophyly. A number of synapomorphies can belisted for this natural entity (Smith and Van Wyk 1991)thus confirming their common origin. It is unlikely thatthe combination of the distinctive alooid karyotype (x=7,four long and three short chromosomes), and thecharacteristic leaf morphology have arisen more thanonce. Furthermore, the vascular bundles of leaves ofAloaceae typically have a well developed cap of thin-walled parenchyma cells at the phloem pole. These areoften referred to as aloin cells (Smith and Van Wyk1992b).

General characteristics

All species of Aloaceae are slow-growing, petaloid,rosulate or distichous perennials. They differ in size fromminiatures barely 10 mm high (Haworthia parksiana) totrees of massive bulk up to 20 m tall (Aloe barberae).Almost all species are long-lived succulents that haveabove-ground storage organs (leaves, stems), but a fewhave subterranean storage and perennating organs (e.g.Chortolirion). Very few are deciduous in the sense thatthey die down and become dormant for part of the year(usually the cold, dry season). Floral morphology isvariable and some basic forms occur repeatedly indifferent genera, which adds another dimension to theinterpretation of the patterns of variation in reproductivemorphology. In general, representatives of the familyAloaceae are easily recognised by their rosettes of usuallyspiny or tuberculate leaves that with age wither frombelow. The adventitious root system is usually shallow andutilises the upper leaf litter layer of the soil. In somespecies the roots may be fusiform and deeper. Theracemose or paniculate inflorescence appears to beaxillary but is in fact borne apically, bearing fleshy tubularflowers. The fruit is a three-angled, oblong, loculicidalwoody capsule. In this regard Lomatophyllum with itsfleshy berries is an exception.

The aloes often form a conspicuous feature of thesouthern African landscape and are popular gardensubjects with their tough, thorny leathery leaves. Mostspecies of Aloe have large, colourful, tubular flowerswhich are pollinated mainly by birds. Flowering is usuallyin the dry season and the seeds ripen just before the rainyseason. Representatives of the other alooid genera areusually cryptic dwarf plants which occur in the shade ofnurse plant species. Haworthia and Astroloba have smallinsect pollinated flowers, whilst bird pollinationpredominates Gasteria and Poellnitzia.

The genera of Aloaceae differ from the closely relatedAsphodelaceae in their conspicuous succulent leaf

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consistency, crescentiform or cymbiform leaf outline incross-section, and the markedly bimodal karyotypeconsisting of 2n=14 chromosomes (Smith 1991a). 3n thisregard Bulbine and Kniphofia appear to be problematic,particularly since some of the species of Bulbine havekaryotypes and morphologies similar to that of certaintaxa of Aloaceae (Rowley 1954). However, Bulbine can beeasily distinguished from genera of the Aloaceae on thebasis of its open, yellow (only very rarely white or orange)flowers, free perianth-segments, bearded filaments, lackof nectar production and the annual nature of some of itsspecies (for example B. alat@. Furthermore, Bulbine hasan African-Australian distribution whereas the Alooideaeis absent from Australia. Mainly for these reasons BuZbinewas not considered to be a constituent of the Aloaceae. Incontrast to representatives of Bulbine and the Aloaceae,leaf succulence is absent in Kn@hofiu, the leaf outline(cross-section) is V-shaped and it has a chromosome basenumber of six.

Aloe occurs over much of sub-Saharan Africa, rangingfrom the southern tip of Africa to the Arabian Peninsula.It is also found on Madagascar and Socotra. The genus isecologically heterogeneous within its range of distributionand has diversified into almost every possible habitat,ranging from deserts, grassland and savanna tocomparatively high rainfall coastal forest types. Thefleshy-fruited LomatophyZZum is restricted to theMascarene Islands. The general distribution patterns ofAstroloba, Gasteria, and Haworthia closely resemble oneanother. These three genera are endemic to southernAfrica and are more or less restricted to the summer-dry,semi-arid coastal regions below the inland escarpment ofthe subcontinent. Gasteria and Haworthia have outliers inthe arid river valleys of Natal, Swaziland, and the easternTransvaal, with a single species of Haworthia CH. venosassp. tesseZZata) occurring in the climatically severe central-southern Africa. The distribution of Astroloba is morerestricted than those of Gasteria and Haworthia and it isusually found in slightly more arid environments of theFynbos and Succulent Karoo Biomes of southern Africa.These three genera and Aloe have relatively largenumbers of species indigenous and endemic to the aridsubtropical transitional thickets of the eastern Capewhere they show signs of active speciation. Of the generaof Alooideae, the monotypic Poellnitzia has the mostrestricted distribution. This genus is found only in theRobertson and Bonnievale districts of the south-westernCape Province. In contrast, the other monotype,Chortolirion, is widely distributed in the summer rainfallgrasslands of southern Africa. The genus does, however,enter the winter rainfall region in southern Namibia.

Most species of Aloaceae use chemical (bitter leafsap) or mechanical (pungent, acuminate leaf tips andprickles) strategies to prevent predation (Van Jaarsveld1987). Many taxa occur in grassland and savanna habitatswhere natural fires are common. In the case ofarborescent species fire adaptation is usually through the

retention of a continuous cover of persistent, dead leavesaround the stems, which acts as an insulator. Smallerspecies are geophytic or semi-geophytic with subterraneanstolons. A few species have true bulbs. Many species ofAloe of the dry Mediterranean Fynbos regions are alsosubject to regular fires and often have resproutingabilities (e.g. the shrubby A. commixta group which havelignotubers). The arborescent A. plicatillis has a corkybark. A. pearsonii of the dry, winter rainfall Richtersveldand Namib regions has functional persistent leaves formost of the height of the plant. These become turgidduring a good rainy season. Species of Gasteria occurmostly in dense vegetation, and with their green, mottledleaves are well camouflaged. When browsed they willresprout from broken leaf fragments. Species ofHaworthia are usually small and frequently occur inspecialised habitats.

The southern African alooid genera

Aloe L. The genus Aloe has been thoroughly treated byReynolds (1966, 1982). A revision of the South Africanspecies has recently been undertaken by Dr. H. F. Glenand Mr. D. S. Hardy. Aloes have tremendoushorticultural appeal in South Africa. The bright flowersand interesting leaf ornamentation, and the horticulturalhardiness of most species ofAloe make them attractive forcultivation even under less favourable hot and dryconditions. Though the majority of species are welladapted to arid environments, few can survive lowtemperatures and frost, and severely cold spells willoccasionally lead to the demise even of individuals ofspecies that occur naturally above the climatically severeinland escarpment (e.g. Aloe greatheadii var. dua~yn~zlr;pers. obs. by G. F. Smith). To many South Africans aloesare what roses and rhododendrons are to Europeans, andthese rosulate succulents will always be horticulturallyimportant in that country. An AZoe craze was sparked offin South Africa in the 1960s and 1970s by the seminalworks of Groenewald (1941) and Reynolds (1966, 1982)resulting in the publication of numerous popular works onthe genus. These include the works of Beyleveld (1973)Bornman and Hardy (1971) Jankowitz (1975), Jeppe(1974, 1977) Judd (1972) and West (1974). In addition,the Aloe Breeders Association is an active group ofamateurs that continuously register new cultivars. Interestin the genus eventually resulted in legislation broughtforward to protect species in their habitats, especially toprevent indiscriminate collecting. Nowadays, as a result oftheir horticultural appeal, most African states imposestrict legislation to protect species of Abe (see section onConservation Measures, Southern Africa, Chapter 3).

Unfortunately, some Aloe habitats have beendestroyed by agricultural activities and urban expansion,and to a lesser extent by collectors and for ethnic uses.Fortunately, some species have restricted habitats notsuited for agricultural purposes, and species have beensuccessfully introduced into cultivation and seeds are

12

freely available. Rarest and most threatened of all is A.polyphylla, the species having novelty and ornamentalappeal. Its numbers have drastically been reduced bycollectors. However, the probability of it becoming extincteven in habitat appears to be slight. Unfortunately, fewspecimens of A. polyphylla survive ex situ due to itsspecialised habitat requirements. Smith (1989) showedthe destruction of the natural habitat of A. bowiea and A.myriacantha in the south-eastern Cape, and emphasisedthe importance of the acquisition of land for itsimmediate protection. A. ferox is of economic importanceand is one of the most common of all species. It is easilycultivated and readily establishes from seed. Commercialplantations of A. ferox have been started at Albertinia,South Africa. Concern has been expressed on the effectthat the removal of the leaves has on survival of thespecies in the wild, since the dried leaves protect it againstfires. The local AZoe sap tappers industry was worth R2.5million in 1992 (Newton 1993).

Haworthia Duval. The genus has been taxonomicallytreated by Bayer (1982) and Scott (1985). The treatmentof Bayer (1982) reflects a more natural dispensation.

Haworthia consists mostly of dwarf rosulatesucculents, often proliferating from the base and formingdense clusters with smooth to tuberculate (rarelypubescent) leaves. The lax racemose inflorescences bearinsignificant, bilabiate, whitish flowers. Of the alooidgenera, Haworthia is under most threat due to habitatdestruction. These usually cryptic plants are oftenconfined to Renosterveld (sclerophyllous shrublands),Succulent Karoo, and arid savanna regions. TheRenosterveld regions are of agricultural importance in thesouthern Cape and most have been transformed to wheatfields, with the result that many populations have beendrastically reduced and a few species could becomeextinct if drastic measures are not implemented. Smith

(1991~) drew attention to the reduction in size ofpopulations of H. fasciata in its south-eastern Capehabitat. Most species have tremendous horticultural valueand because of their small size and relative ease ofcultivation are popular container plants and highly prized,especially in Japan, Europe, and the USA. Somecollectors have also played a role in reducing numbers.This is, however, minimal in comparison to agriculturaldamage to their habitats. Under the leadership of Mr.Kobus Venter, two Haworthia localities (H. magnifka var.maraisii and H. mirabilis var. badia) within the Napiervillage municipal region have been declared nationalheritage sites. Furthermore, many species of Haworthiaare being multiplied by seeds and leaf cuttings byinteres ted members of the Succulent Society of SouthAfrica. H. ZimifoZia is used medicinally by various Africantribes and its numbers have been reduced considerably inthe wild. However, this species is cultivated at theSilverglen Nursery in Durban and plants are supplied ataffordable prices to traditional healers. Other threatenedspecies in the southern Cape include H. emeZyae var.multifolia, H. magnifica var. atrofusca, H. serrata, and H.poellnitziana.

Gasteria Duval. The genus has been revised (VanJaarsveld 1991) and a more complete account is found inVan Jaarsveld (1994). It is a well defined, monophyleticgroup with brittle, mottled, keeled leaves which are oftentuberculate and usually without spines. It has pendulouspedicels and gasteriform, trichromatic, curved flowers.The black, winged seeds are wind dispersed. The genushas centres of endemism in arid savanna regions and to alesser extent in the Succulent Karoo of the south-easternCape. All species have horticultural potential and they arepopular container plants. A number of species areconfined to the fertile inter-montane valleys of theeastern Cape which is overgrown with subtropical thicketvegetation or grassland or, to a lesser extent, renosterveld.This region is of agricultural importance and large areashave been cleared for crop farming. This region is thehabitat of G. bicolor var. bicolor, G. nitida var. nitida, G.nitida var. armstrongii, G. pulchra, and G. excelsa. Ofthese, G. nitida var. armstrongii is endemic to the lowerGamtoos River Valley and is under threat. Fortunatelythe species is represented in a nearby nature reserve. G.bicolor, although still common, has been reducedconsiderably by agricultural development. G. excelsaoccurs on forested, well-drained rocky banks, but due toovergrazing and firewood collecting has become muchrarer. G. baylissiana is another very rare species from thesummit of the Zuurberg. The reason for its in situ declineis unknown but the numbers of the species have recentlybeen increased by reintroductions to its habitat. Stocks ofthe species have been increased ex situ by seed andvarious clones originally distributed by Col. R. D. Baylissto botanical gardens were used in the project. G. croucherifrom Natal has became rare due to medicinal and ethnic

13

uses in that region. Seed of this Gasteria have been madeavailable to Silverglen Nursery, Durban, whence plantsare provided to traditional doctors. Fortunately, theremaining species occur in habitats not under immediatethreat. All species of Gasteria are being increased fromseed as well as vegetatively at the Kirstenbosch BotanicalGarden, especially the rare endemics, and seeds are madeavailable on an annual basis.

AstroZoba Mitewaal. The genus is closely related toHaworthia, but for some reason has never appealed tocollectors and species of this genus are not underimmediate threat.

Poellnitzia Mitewaal. At present urbanisation does notpose a threat to populations of Poellnitzia rubriflora. In theentire Succulent Karoo Biome it occurs in only twomedium-sized towns (Worcester and Oudtshoorn) andvarious smaller centres, for example Robertson andBonnievale. Cultivation of especially wine grapes islimited to fairly narrow strips alongside water courses,such as the Bree River. The colonies of P. rubriflora aremostly confined to rocky hillocks and appear to be in noimmediate danger of agricultural activities. However, dueto its small world population and its confinement to arestricted geographical area, Poellnitzia has been removedfrom the Indeterminate (I) category designated to it byHall et al. (1980) (under the name Haworthia rubriflora inHall and Veldhuis (1985) and reclassified as Rare (R) inHilton-Taylor (2996b) (see Annex 16 for definitions of theRed List categories).

Chortolirion A. Berger. The genus has a very effectivedefence mechanism in that, in its natural grasslandhabitat, casual succulent plant collectors are likely tomistake its linear leaves with those of a grass species. Theinflorescence and flowers of Chortolirion are also veryinconspicuous. In pristine and well-managed pasturesgrazing pressure is minimal since the abundance ofpalatable grass species associated with it will certainly beutilised preferentially. Therefore, at least at present,Chortolirion is Not threatened. Where possible,populations should be monitored for decline orexpansion, especially since Chortolirion occurs in some ofthe most densely populated areas of South Africa.

AsclepiadaceaeFocke Albers and Ulrich Meve*

The family Asclepiadaceae R.Br. is characterised by itsunique complicated flower morphology and highlyevolved pollination system. The family comprises variouslifeforms including herbs, shrubs, twiners, succulents, andtrees. The species mainly occur in the subtropics andtropics around the world, and only occasionally intemperate regions.

* Manuscript submitted in 1993

Economically the family is of little value. Some generaare of horticultural interest, but most species are not ofsignificant value in terms of horticultural production.Asclepias, Hoyas and related species are grown asoutdoor or pot plants, Dischidias, stapeliads, andCeropegia species can be found all over the world inspecial hobbyist collections. Other uses of the family arefew. The hairs of Asclepias and Calotropis seeds weresometimes used as a substitute for down, and in Indiaropes are produced from the stem fibres. In Africa andIndia some species are used medicinally by local people.Species of Caralluma and Ceropegia are eaten in India(see Annex 10) and, in extreme dry areas of southernAfrica, succulents are eaten by Nama people andBushmen.


Brummit (1992) compiled 315 genera in this large family.Today the number of species is estimated at more than2000. In an intensive study of the family, Liede and Albers(1994) determined that about 500 correctly describedgenera exist. These are arranged in taxonomic order anddiscussed below. The family is divided into threesubfamilies (Periplocoideae, Secamonoideae,Asclepiadoideae), the latter one comprises four tribes(Marsdenieae, Asclepiadeae, Gonolobeae, Stapelieae)according to Bruyns and Forster (1991). A fifth tribeFockeae is added to this subfamily by Kunze, Meve andLiede (1994).

The subfamily Periplocoideae is a well-defined taxon,which is sometimes excluded from the Asclepiadaceaeand treated as a family of its own (Periplocaceae)(Schlechter 1905; Bullock 1956). Most of the species areshrubs and twiners occurring in the Old World. Only twoof the described genera, Raphionacme and Petopeda,have fleshy root stocks. Raphionacme is becoming ofincreasing interest for the hobbyist growers. Venter andVerhoeven (Bloemfontein, RSA) have contributedseveral valuable papers on this taxon since 1983 andGoyder (Kew) is also dealing with this group.

The subfamily Secamonoideae (five genera) isdominated by the great palaeotropic genus Secamone(Australia, Forster and Harold 1989; Africa, Goyder 1992;Madagascar region, Klackenberg 1992). Many species areknown only from one or a few specimens, and 84 per centare endemic to Madagascar (Klackenberg 1992). Thetwiners and herbs are not important in cultivation.

The largest subfamily is the Asclepiadoideae. Withinthis grouping, the tribe Fockeae contains three generawhich are excluded from the tribe Marsdenieae. Thesmall African genus Fockea consists of well-knowncaudiciform succulents. Research on this genus has beencarried out by Court (Grahamstown, RSA). One species,Fockea sinuata, is considered to be Endangered. Ingeneral, the Fockeae species are of no importance foramateur growers.

The tribe Marsdenieae contains attractive leaf

14

succulents. Many of these are epiphytes of East Asia,Australia, and Oceania: Dischidia, Dischidiopsis, andHoya. Some common species of these genera are grown ashouse plants, but most of the species in collections arecuttings of plants of natural origin, which are traded by afew professional horticulturists. Rain forest species in thecentres of diversity in Malaysia, Indonesia, Philippines,and New Guinea are extremely threatened by forestdestruction. Research is being carried out by de Koning(Leiden) and H. and R. Donkelaar (Werkendam), bothfrom the Netherlands, and by Forster and Liddle, fromAustralia.

The largest tribe, Asclepiadeae, has a world-widedistribution in temperate, subtropical, and tropicalregions. Lifeforms represented include herbs, shrubs,twiners, and trees, with succulents relatively uncommon.Large genera o f th i s t r ibe a re A s c l e p i a s ,CynanchumlVincetoxicum, and Oxypetalum. The firstgenus is better known (e.g. Woodson 1954) whereas theothers have been neglected for a long time. Cynanchumand related groups are being studied now by Liede (Ulm,Germany) and Nicolas (South Africa). The species of thegenera which occur in the New World are quite differentfrom those living in the Old World.

Succulents in the Asclepiadeae include species ofCynanchum with fleshy leaves, and species of Asclepias,Aspidonepsis, Cynanchum, Sta tmostelma, and Stenostelmawith succulent subterranean parts. Stem succulentsinclude species of Sarcostemma, Cynanchum, Folotsia,Karimbolea, and Platykeleba, all of which, with theexception of Sarcostemma, exist in the semi-arid parts ofsouth-west Madagascar. Some of these are extremelylocalised or rare. Liede (pers. comm.) listed 48 species ofCynanchum, five of Folotsia, two of Karimbolea, andPlatykeleba insigne as Rare and Endangered succulentMalagasy species. Several of them have apparentlydeclined rapidly in numbers as their habitat is graduallybeing destroyed. The stem-succulents are of interest tohobbyists, but other species are not considered desirableand therefore they have not been additionally endangeredby professional collectors.

The species of the tribe Gonolobeae are restricted tothe warmer parts of Central and South America. Theherbaceous species often belong to monotypic generawhich are poorly known in the wild and in herbariumcollections. Several species are represented by singleherbarium sheets and some species have never been re-collected since they were first described. Stevens(Missouri Botanic Garden) is working on this verydifficult and poorly known group.

The Stapelieae is by far the best-known tribe and themost important one for hobbyist growers. All species areof palaeotropical origin, and many have evolved lifeformswith fleshy subterranean parts or stem succulence. Thetribe Stapelieae includes the genus Ceropegia and a groupof 30 other genera which collectively are commonlyknown as stapeliads.

The distribution area of Ceropegia covers Africa,Madagascar, Arabia, India, Southeast Asia, theIndonesian Archipelago, and Northern Australia.Although the genus is so widespread, most of theapproximately 200 species are endemics with restricteddistributions. Due to this fact, and the horticulturalappeal of Ceropegia spp., the genus as a whole is listed inAppendix II of CITES. Within the Stapelieae, this genushas the highest rate of insufficiently known species (20 percent). Since Huber’s revision of the genus in 1957, severalnew species have been described which are highlyattractive to hobbyists. Most, except for the geophytcs,can be easily propagated by cuttings, and have been incultivation for many years.

The species of the genus Brachystelma (about 120) aswell as those of Riocreuxia and Tenaris, all have developedgeophytic forms and are well-adapted to grassland. Thespecies of Brachystelma have a similar distribution area asthose of Ceropegia to which they are closely related, butthey avoid wetter areas. The habitats of Bnrchystelma areunder extreme grazing pressure. The species do not growwell on their tubers in cultivation, and can bc propagatedonly by seed. According to the species listed in the“Checklist of Brachystelma, Ceropegia, Riocwuxia, and theStapelieae” (Boele, Kroesen, and Noltee 1987) 72 percent of the approximately 120 species are Endangered. Sixspecies are already Extinct or have not been found for along time. These are B. gemmeum, B. glcwse, B.gracillium, B. longifolium, B. na talense, and B. occidefz tale.Valuable information is given by Dyer (1983). The genusMacropetalum is insufficiently known.

The thirty genera of stapeliads contain approximately400 species. As stem succulents, with often large andattractive flowers, they are very well known to hobbyistgrowers and can be found in collections all over the world.In the wild, most of the species are confined to the morearid parts of Africa, but there are also several speciesoccurring in Arabia, India, and Myanmar. Two Europeanspecies are closely related to those of North Africa. The

Orbeopsis gerstneri ssp. gerstneri, a South Africanasclepiad.

15

species mainly form small, widely scattered or sometimessolitary populations. It is becoming increasingly apparentthat stapeliads are a comparatively old plant group, inwhich plate tectonics and changes of climate have led toseveral migratory processes. The present distributionpattern comprises not only various centres of diversity,but also very restricted areas of retreat (Albers and Meve,in prep.). This perspective allows for the introduction of arelatively large number of genera in the stapeliads, a viewsupported by most of the recent studies. Research work sofar, however, has mainly been limited to morphologicalcharacters and unfortunately has led to divergentconcepts of species and even genera. Experts are far froman acceptable agreement like the one reached by the 10sconsensus group for the Cactaceae.

Northeast Africa is assumed to be the centre ofevolution for the stapeliads, but most of the species occurin southern Africa today. This secondary centre (Albers1983) has evolved its own genera: Hoodia, Huerniopsis,Lavrania, Notechidnopsis, Ophionella, Orbeanthus,Orbeopsis, Pectinaria, Quaqua, Stapelia, Stapeliopsis,Tavaresia, Trichocaulon, Tridentea, and Tromotriche. Thegenus Stapelianthus is endemic to Madagascar. Only a fewgenera, like Huernia and Duvalia, are known on both sidesof the equator, the vast majority of the North Africanspecies belongs to the genus Caralluma. The species fromIndia and Myanmar are included here. Despite intensiveresearch which has been done by Gilbert (1990) the genusstill needs attention. With the aid of modern methods itseems to be possible to elucidate this complex (Albers andco-workers, in prep.).

Most of the other large genera of the stapeliads havebeen revised recently or revisions are in preparation(Duvalia, 14 spp., Meve, in press; Echidnopsis, 31 spp.,Bruyns 1988; Hoodia, 13 spp., Bruyns 1993; Huernia, 69spp., Leach 1988; Lavrania, 5 spp., Bruyns 1993; Orbea, 21spp., Leach 1978; Orbeopsis, 11 spp., Leach 1978;Pachycymbium sensu Gilbert, 31 spp., Gilbert 1990;Piaranthus 7 spp., Meve 1994; Quaqua 24 ssp., Bruyns1983; Stapelia, 45 spp., Leach 1985; Tridentea, 20 ssp.,Leach 1980). Valuable information on the distributionand ecology of the species is included in most of therevisions.

The HoodialTrichocaulon complex, which has alsobeen studied by Plowes (1992) still needs a satisfactorynomenclature. Lavranos has contributed to the knowledgeof most of the taxa in a valuable way by describing newspecies and introducing new genera which he had foundand collected on field trips in Africa and Arabia (mainlypublished in the Cactus and Succulent Journal (US).

Little is known about the population behaviour of thestapeliads. Observations on Hoodia spp. in Namibiaindicated that some specimens have a lifespan ofapproximately 60 years (Jurgens ex Erb, pers. comm.).Albers and Meve (1991) demonstrated, in contrast, thatsome karroid species with creeping lifeform behave] likeephemerals, e.g. within nine years a large Duvalia

caespitosa population wa s subst ituted by a Stap eliapillansii population. Fur ther lo ng-term studies ) ofenormous importance for studying conservation status,are needed for other taxa.

The myiophilous pollination syndrome of thestapeliads is now being investigated more carefully (Meveand Liede 1994). The knowledge of pollinators of theother taxa is still very poor. A literature review ispresented by (Meve and Liede 1994).

Species of the stapeliads can be quite readilypropagated by seed, but most of the specimens which arccultivated by specialists are of wild origin or from materialexchanged among colleagues. Many of the species are noteasy to grow on their own roots, so grafting seems to be avery successful way to keep them alive and make thembloom sooner.

Threats

The destruction of the natural habitat is by far the mostthreatening danger for the species. In the winter rainfallarea of South Africa, as well as in East and North Africa,overgrazing and the spreading of agriculture is the mostimportant cause of this destruction. The karroid regionsof South Africa seem to be rather stabilised, whereassome Fynbos species are known to be threatened byagricultural activities and housing development on thecoast. Species threatened in this way include those ofDuvalia, Orbea, and Stapelia. The problems facingsucculents of the arid regions of Madagascar aredescribed in the Madagascar account in Chapter 3. Mostof the Stapelianthus spp. of Madagascar are underenormous pressure except S. decaryi and s.madagascariensis. The Indian Caralluma species are allconsidered to be threatened due to spreading agricultureand population pressure. The monotypic Frerea indica ofIndia (often only accepted as Caralluma frerei), togetherwith the genus Ceropegia, are the only stapeliads listed inCITES (Appendix II). Caralluma europaea is threatenedby the increased building of holiday villages in Spain. The

Stapelianthus pilosus

16

endemic C. hurchardii of the Canary Islands seems to besufficiently protected. This species, the two EuropeanCarallumas and the Ceropegias of the Canaries areprotected by EC legislation.

Although the field collecting of amateurs should notbe underestimated, as a threat it is secondary to habitatdestruction. It is controlled by governmental restrictionsin several countries.

Conservation status

Using the updated “Checklist” (Boele, Kroesen, andNoltee 1987) which is mentioned already, Meve (pers.comm.) compiled all data available for the Stapeliads.Following the old IUCN Red List categories (see Annex16) he found out that no species is Extinct, but that 12.6per cent of the species are Endangered, 1.8 per cent areVulnerable, 29.5 per cent are Rare, and 7.5 per cent areIndeterminate. That means that half of the Stapeliadspecies are threatened today. The same percentage isfound with Ceropegia if the rather high number ofinsufficiently known species is included. A list of theAsclepiadaceae with conservation categories is given inAnnex 2.

To summarise, the Asclepiadaceae comprises morethan 2000 species with most occurring in the subtropicsand tropics. The more arid, as well as the wetter areas ofthese zones, contain very sensitive ecosystems which areunder considerable pressure or have already beendestroyed. Many species of Asclepiadaceae are thereforethreatened with extinction, including many species whichare poorly known. Long-term studies are needed nearlyeverywhere. Within the best-studied tribe Stapelieae, halfthe species are considered to be threatened. In othergroups the percentage is even higher. In general, thereasons affecting the conserva t ion s ta tus o fAsclepiadaceae are increasing agricultural development,forest destruction, housing, and road development.

CactaceaeNigel P. Taylor

The Cactus family (Cactaceae Juss.) is characterisedbotanically by the presence of stems bearing specialised,felted short-shoots termed areoles, which usually developthe spines (modified leaves) that represent the familiarhallmark of this plant group. Except for one species ofRhipsalis (R. baccij’era - Neotropics eastwards to SriLanka), the family is endemic to the New World, althoughvarious economically important and weedy species havebeen introduced and become naturalised in warmer partsof the Old World. In the Americas they range fromsouthern Canada to Patagonia, being most frequent in thedry climatic zones between 35” N and S latitude andoutside the moist equatorial (Amazonian) region. Thefamily has very considerable value in terms ofhorticultural production, with millions of artificially

propagatedeach year.

Taken awide clima

specimens traded across international borders

s a whole the Cat tus family has a surprisinglytic and ecologica.l spectrum, encompassing

almost rainless desert such as parts of the westernAtacama in northern Chile at one extreme to tropical rainforest receiving more than 2000 mm of rain per year atthe other. They range from sea level to a reported 5200 maltitude in the Andes and vary considerably in theirresistance to frost. None is parasitic, but epiphytes,lithophytes, ‘cactus geophytes’, mound-formingcaespitose, decumbent, scandent and erect, free-standingtree and shrub species are frequent. Apart from the‘cactus geophytes’, amongst the most remarkable arcdwarf species restricted to cliffs, such as Aztekirrm,Strombocactus, and Blossfeldia, which have dustlikestrophiolate seeds. In contrast, Opurztia (Bi.(~,~ilio~)Lt~~ti~~)brusiliensis is adapted to life in high, dense, dry to veryhumid forest, where it forms a tree to 20 r-n, with acylindrical trunk and flattened leaflike ultimate stem-segments (southern Neotropics). At least four Mexicanspecies appear to be restricted to gypsum, and isolatedoutcrops of particular rock types account for a significantpart of the narrow endemism seen in the family. Somespecies are able to grow on, or are restricted to, extremelyoligotrophic substrates, for example, Uebelmamlia andDiscocactus spp. Seed dispersal is effected by variousmeans including mammals, birds, lizards, insects(especially ants), fish (Amazonia), wind (Eriosyce spp.,Pterocactus), and water (Frailea). Bats are a particularlyimportant group of associated organisms (Dobat andPeikert-Holle 1985) on whose conservation the survival ofsome cactus species may well depend. Nurse plants ofvarious shapes and sizes are often important forestablishment of delicate cactus seedlings (Cactoideae).

Although amateur horticulturists continue to play asignificant role in describing and classifying cacti, the past20 years have seen the family subjected to fairly intense

Azfekhm hintonii, Nuevo Leh, Mexico.

17

morphological and taxonomic scrutiny by professionalbotanists and it is now probably one of the betterunderstood major succulent plant groups. The Cactaceaecomprises four subfamilies, namely the primitively leaf-bearing Maihuenioideae and Pereskioideae (2 genera / c.19 species), the highly derived Opuntioideae (5 or moregenera / 185-265 spp.) with barbed spines and glochids,and the leafless, morphologically complex Cactoideae (c.97 genera / > 1000-2000 spp.). The last of these,containing the bulk of the family, is divisible into at leasteight tribes, each of which has a characteristicgeographical range.

Collaborative study of the family’s classification atgeneric level by a broad group of botanists andknowledgeable amateurs (under the auspices of the 10s)has resulted in two published reports listing the generaaccepted by a consensus of systematists (Hunt and Taylor1986, 1990). Following these reports, a checklist of speciesnames and their synonyms in current usage has beencompiled for the family on a consultative basis. Thisincludes distribution data at country level and country bycountry species lists with details of endemism (Hunt1992). This checklist, prepared at the behest of the CITESSecretariat, distinguishes between species fully accepted assuch according to certain primary sources (botanicalmonographs, floristic treatments, and in the opinions ofauthors of a few unpublished taxonomic revisions) andthose only provisionally accepted, which include taxawhose status awaits expert botanical study and also thosetreated as subspecies or geographical varieties in theprimary sources consulted. In total, the Cactus familycomprises about 105 genera and, following the CITESChecklist, 1208 accepted and a further 1300 provisionallyaccepted species. If infraspecific taxa are excluded fromthe latter figure, leaving those whose taxonomic status isleast understood, then it becomes clear that the greatestconcentrations of taxonomically doubtful species arefound in Peru and Bolivia. It is this region of the centralAndes where competent field study is most needed toresolve questions of taxonomic and conservation status.Nevertheless, it is probable that a significant proportionof suchso their

taxa willnumbers

prove to be ‘good’ endemic species andshould be ta.ken into consideration when

analysing this Andean region from a conservationperspective.


In spite of the above-mentioned uncertainties, it is arelatively simple matter to identify the geographicalcentres of diversitv and of endemism for cacti. In firstplace by far comes Mexico and the immediately adjacentsouth-w est ern United S tates (southern California,Arizona, New Mexico, and south-west Texas) where some27 per cent of all cactus genera are endemic and at least570 accepted species are native, 430 being endemic toMexico alone (27 cactus species have been reportedsympatric in a single locality in north-east Mexico, Fitz

Maurice, pers. comm.). The region is notable for the highnumber of endemic tree-like or columnar speciesbelonging to tribe Pachycereeae, which are characteristicof the dry tropical forest and Sonoran Desert floras,where they sometimes dominate or are so conspicuous asto be worthy of description as forming ‘cactus forest’.Many are of local economic importance for their ediblefruits, woody skeletons used in house construction, andfor planting as impenetrable living fences. Bats are knownto be important pollinators and seed dispersers of thesecolumnar cacti. Mexico and the bordering Rio Grandevalley region of the USA (especially the ChihuahuanDesert and its margins) have many remarkable small ormonotypic genera from tribe Cacteae, which exhibitunique and sometimes extraordinary morphology, forexample, Ariocarpus (7 “pp.), Astrophytum (4 “pp.),Aztekium (2 spp.), Epithelantha (2 “pp.), Geohintonia (1sp.), Leuchtenbergia (1 sp.), Lophophora (2 spp.),Mammilloydia (1 sp.), Obregonia (1 sp.), Ortegocactus (1sp.), Pelecyphora (2 spp.), Stenocactus (8 spp.), andTurbinicarpus (14 spp.). Most of these include specieswhich are either of very restricted distribution or foundonly in very unusual habitats, and some are known to beunder threat from collecting for horticultural purposes,which has required their protection from commercialover-exploitation by inclusion in Appendix I of CITES.Other low-growing genera, which are particularly diverseand species rich in the south-west USA and Mexico,include Mammillaria (> 150 spp.), Coryphantha (> 50spp.), and Echinocereus (> 50 spp.), each with threatenedtaxa, some of which are included in Appendix I of CITESin view of horticultural demand. Apart from collectingpressure the most threatened taxa are those ‘cactusgeophytes’ that inhabit ferti le land undergoingagricultural development (e.g. Echinocereus prrlchellus).Opuntioideae, i.e. Opuntia spp., make up a very importantcomponent of this cactus flora and some are ofconsiderable local economic importance for their ediblefruits and stem-segments. Southern Mexico has asignificant number of species belonging to the primarilyepiphytic tribe Hylocereeae (e.g. Disocactus s.l.),inhabiting moister tropical forest and mountainousregions, but in comparison with tribes Cacteae andPachycereeae, a smaller proportion of these is endemic,with ranges frequently extending into the adjacentcountries of Central America, where they represent themost important component of the cactus flora.

The second most diverse region for cacti is that of thecentral Andes comprising the countries of Peru andBolivia, with the addition of southern Ecuador, north-eastChile, and north-west Argentina. About 18 per cent of allcactus genera are endemic to this region and there areabout 420 taxa named as species endemic to Peru andBolivia, but most of these are only provisionahy acceptedby the CITES Checklist. The cacti of southern Ecuador,north-east Chile, and north-west Argentina are betterknown and comprise about 115 accepted specie s. As in the

18

case of Mexico, cactus forest vegetation with large shrubsand columnar or tree-like forms occurs, but highlyspecialised and unique growth habits are less common,exceptions being represented by Blos@eZ&a (1 sp.) and anunusually diverse array of Opuntia spp. However, thereare numerous species of globose habit from tribesNotocacteae and Trichocereeae. Genera containing manyspecies include Parodia, Rebutia, Gymnocalycium,Echinopsis, Cleistocactus, Oreocereus, and Matucana. Thecommonest pollination syndromes in this region are thosefor bees, hawkmoths, and hummingbirds.

The third most diverse region is that here looselydescribed as eastern Brazil, comprising the segment ofthat great country east of a line drawn between the statesof Maranhao and Sao Paulo (north-east and south-eastBrazil plus eastern Goias and eastern Tocantins). About80 per cent of its cactus flora is endemic, including 11 percent of all cactus genera and having a total ofapproximately 145 taxonomically acceptable nativespecies (plus numerous taxa recognisable as subspecies).Its cacti are perhaps the best understood after those ofthe USA and northern Mexico (Chihuahuan and SonoranDeserts). Columnar, shrubby and sometimes tree-likeforms (tribe Cereeae and Pereskia) and epiphytic forms(tribe Rhipsalideae) predominate, the former beingparticularly diverse in the dry interiors of Bahia andMinas Gerais states, the latter characteristic of the coastalMata Atlantica and mountains of Rio de Janeiro and SaoPaulo (and also the states of southern Brazil). Much ofthe Mata Atlantica has now been destroyed and with itthe habitat of these epiphytic cacti. Rhipsalis pentaptera,for example, although common in cultivation, is probablyExtinct in the Wild, since its only known habitat formerlyoccurred in what is now a built-up area within the city ofRio de Janeiro. The east Brazilian cereoid cacti arenotable for the frequent development of diverse kinds of

woolly lateral cephalia, which in some cases are thoughtto be adaptations connected with pollination bY bats, thecommonest cactus pollen vector in the region.Hummingbird and hawkmoth pollination syndromes arealso frequent. The fewer species of globose habit includethe taxonomically isolated Uebelmannia (3 spp. restrictedto a small area in Minas Gerais state) and Melocactus ( 15spp.) and Discocactus (6 “pp.), with flowers borne inpeculiar inflorescence structures called terminal cephalia.Representatives of each of these three genera arcregarded as severely threatened and are currently the onlySouth American cacti listed in Appendix I of CITES(Braun and Esteves Pereira 1988; Taylor 1992). Membersof Opuntioideae are diverse at generic level, butrepresented by very few species when compared withother areas within the family’s range.

The geographical region that is fourth in order ofimportance comprises central-western and southernBrazil, Paraguay, Uruguay, and Argentina (excluding thenorth-west and southern parts). There are approximately85 accepted species (about half of these endemic) and agreater number of endemic but only provision ally acceptedtaxa. Columnar, shrubby, or sem i-scandent cacti aremainly represented by species from the genera Cerezls,Harrisia, Opuntia, and Pereskia, most of which areendemic, but the greater representation of the family is inthe form of low-growing or globular members of Parodias.1. (incl. Notocactus etc.), Frailea, and Gymnocalyciunz,each with numerous named species and in need oftaxonomic revision. Habitats of various of these smallercacti are shrinking due to agricultural development andfield studies are required to determine their conservationand taxonomic status. There is also a high number ofepiphytic species from tribe Rhipsalideae, whose habitatis shrinking through deforestation, as in eastern Brazil.

Smaller, but nevertheless important, centres of

savanna in Haiti.

endemism include the Caribbean region with northernSouth America (north Venezuela, north and westColombia and Panama), where the most notable generawith endemic species are Melocactus with 9 and Cereuswith 5, and central to north-western Chile (especially thewestern edges of the Atacama Desert), where an isolatedflora has developed. The Caribbean islands have relativelysmall cactus floras, exceptions being Cuba andHispaniola, but are notable for peculiar endemics such asthe Consolea group of Opuntia, Leptocereus (including thegiant Neoabbottia), Harrisia subg. Harrisia, the massiveDendrocereus (now included in Acanthocereus by someauthors), and a unique group of dioecious Pereskia spp.,some of which are very rare and of particular conservationconcern (Leuenberger 1986).

If the north-easternmost corner of Chile is excluded(see above), the remainder of the country has an almostexclusively endemic cactus flora of about 60 acceptedspecies. The coastal fringe of the Atacama Desert innorthern Chile is home to many remarkable cacti whoseexistence depends to a large degree on moisture derivedfrom fogs which roll in from the cold Pacific Ocean.Various remarkable life forms have evolved in this region,including extreme ‘cactus geophytes’ and certain membersof the endemic genus Copiapoa (about 25 spp.) with stemscovered in dense white or grey wax. Excepting a fewhummingbird-syndrome species, nearly all the endemicChilean cacti are thought to have flowers adapted forpollination by hymenoptera. Cacti found in the regions ofSantiago, Valparaiso, and southward include speciesunder threat from urban and agricultural expansion, andat least one member of the largest Chilean genus, Eriosyce(27 spp. endemic), is now believed to be Extinct in theWild (E. aspillagae; see Kattermann 1994).

Towards the extremities of the family’s range, andnear its equatorial centre, few species are represented dueto obvious climatic constraints. In the north, in the USA,should be mentioned the Rare, narrowly endemic speciesof Pediocactus and Sclerocactus, some of which areprotected by CITES Appendix I listing. To the south, inArgentinian Patagonia and adjacent Chile, there arepeculiar representatives of the Maihuenioideae(Maihuenia, 2 “pp.), Opuntioideae (Pterocactus spp.), andtribe Notocacteae (Austrocactus spp.), but none of these isregarded as particularly threatened at present. In theconstantly humid Amazonian region there are only a fewwidespread epiphytic species, plus some poorly knownendemics (MeZocactus and Cereus “pp.), the most notablecactus being Selenicereus wittii, which inhabits floodedforest (igapo), climbing up the trunks of trees by means ofaerial roots (Mee 1988). Farther west, in the Galapagosarchipelago, is a small but entirely endemic cactus floracomprising the monotypic genera Jasminocereus (a closerelative of Armatocereus from the South Americanmainland) and Brachycereus, and various unusual OpuntiaSPP*

Threats

The major pressures affecting the conservation status ofcacti are, in decreasing order of severity, (I) agriculturaldevelopment and deforestation, (2) urbanisation andinfrastructural development including roadbuilding/widening and hydroelectric dam projects, (3)collection for horticulture, (4) mining of stone forconstruction (especially threatening to species restrictedto limestone or granitic rock outcrops, for example, ineastern Brazil and Mexico).

CrassulaceaeHenk t’Hart

The Stonecrop family (Crassulaceae DC.) comprisesherbaceous, and sometimes shrubby, leaf succulents withregular, hermaphrodite flowers which have an equalnumber (commonly five) of sepals, petals, and carpels.With few exceptions, the carpels are many-seeded andcompletely free. The distribution of the family is nearlycosmopolitan, but most taxa are restricted to thesubtropical and temperate regions of the Northernhemisphere and southern Africa. The family is very rarein Australia and South America. The Crassulaceaecomprises a large number of species of great horticulturalvalue, most of which can be easily propagated and areproduced commercially.

As with other succulent plant groups, the taxonomy ofthe Crassulaceae is hampered both by the difference inthe appearance of herbarium specimens compared to thatof live plants, and by a lack of clear-cut discontinuities inmorphological features. The latter is probably due tostrong morphological constraints resulting from theirhighly specialised habit. Consequently botanical opinionstrongly diverges with respect to the number of speciesand genera in the family as well as to the delimitation ofinfrageneric and infrafamilial taxa. The use of living plantmaterial in systematic studies may to a large extent helpto solve these disputes (as demonstrated, for example, byPraeger 1921, 1932; Froderstrom 1930-1936; Walther1972), but at the higher taxonomic levels, especiallyconcerning the delimitation of genera, there is still muchdisagreement among professional botanists. Recently ajoint study by a small group of Crassulaceae experts,under the auspices of IOS, has resulted in the first draft ofa list of genera accepted by consensus (Eggli et al. 1994).

Crassulaceae occurring in regions which have beenextensively explored botanically are fairly well-known.The exact number of species within this family is,however, still uncertain, as knowledge is incomplete forCentral and South America, the Near East, southernCentral Asia (the Himalayas), East Africa, andMadagascar. Despite these gaps in our knowledge, theevolution and systematics of the family are now beginningto be unders tood . Combined cytological andmorphological studies of plants cultivated under uniformconditions have greatly improved our understanding ofthe significance of infraspecific variation and speciation in

20

the family and has considerably improved speciesdelimitation (for references see t’Hart 1991). RecentDNA studies, however, have most of all enhanced ourunderstanding of the evolutionary relationships within thefamily and have helped to verify earlier tentativelyproposed evolutionary trends (van Ham 1994).

The Crassulaceae family comprises 1200-1500 species.In the most recent monograph covering the whole familyBerger (1930) distinguished 6 subfamilies. However, thefamily comprises only 2 major evolutionary lines andconsequently the number of subfamilies has been reducedaccordingly (van Ham 1994; t’Hart 1994). The subfamiliesare the Crassuloideae (sensu Berger 1930), which isdistinguished from all other Crassulaceae by acombination of haplostemonous flowers and oppositeleaves (2 genera / c. 200 species), and the Sedoideaew h i c h i s charac te r i sed by predominantlyobdiplostemonous flowers and spiral phyllotaxis (c. 30genera / 1000-1200 species).

The family probably evolved about 100-60 millionyears ago in Eastern Africa or the Mediterranean region(van Ham 1994). From this primeval centre of origin twolineages, the Crassuloideae, and the Cotyledon andKalanchoe lineage of the Sedoideae, migrated intosouthern Africa, whereas the other lineages of theSedoideae spread over Europe, Asia, and North andCentral America in four or more successive migrationwaves.


With the exception of the large genus Sedum (about 500species), which is widely distributed in the subtropical andtemperate regions of the Northern Hemisphere, and thesemi-aquatic, cosmopolitan genus Tillaea (about 20species), the genera of Crassulaceae are generallyconfined to a single continent or subcontinent. Four main

geographical centres of diversity and of endemism can bereadily distinguished, one in southern Africa and three inthe Northern Hemisphere. These four centres differlargely with respect to the species and the genera theycomprise. The three centres in the Northern Hemisphere,do however, each contain a unique section of Sedzrm(t’Hart 1982).

The southern Africa centre of diversity hasapproximately 450 species in six major genera, C~uss14la,Adromischus, Cotyledon, Bryophyllum, Kulanchoe, andTylecodon. Altogether they constitute the larger parts ofthe subfamily Crassuloideae and the entire Kulanchoe andCotyledon lineage of the subfamily Sedoideae (t’Hart1994) which, with few exceptions, are all characterised byopposite leaves. South Africa is by far the richest regionwith over 140 species of Crassula (Tolken 1977) and mostof the species of Adromischus, Cotyledon, and Tylecodon.Crassula s.1. comprises some highly specialised,systematically isolated forms which are sometimes treatedas separate genera (small or monotypic), e.g. Dinacriu,Pagella, Rochea, and Vauanthes. Crassula also occurs intropical East Africa and Madagascar, but it isoutnumbered there by Kalanchoe (over 100 spp.).Bryophyllum (incl. Kitchingia) is endemic to Madagascar.The systematics of the East African Kulunchoe species hasrecently been reviewed in a series of local Floras, but theMadagascan species of Kalanchoe and Bvop/1yll14rz1, mostof which are probably endangered, are urgently in need ofrevision. They show a remarkable series of physiologicaland morphological adaptations to different habitats andpollinators. Some Bryophyllum species producebufadienolides, (cyto)toxic secondary compounds whichare regarded as potent, novel antitumor agents(Yamagishi et al. 1989).

In East Africa, from Madagascar to Ethiopia, some 15rare and aberrant Sedoideae of uncertain affinities arefound. These include Afrov ive l la , Crassrrlnriu,Hypagophytum, and Perrierosedum, and may be relicts ofan ancient (Tertiary), African Sedum flora.

The second centre of diversity is central Mexico withabout 300 endemic species in five major genera(Echeveria, Graptopetalum, Pachyphytl4m, Sedum,Villadia). The states of Puebla, Hidalgo, and Oaxaca areextremely rich in species, containing over 60 species ofEcheveria alone (Walther 1972). This region adjacent toand including the eastern section of the TransmexicanVolcanic belt also comprises a wide variety of otherCrassulaceae. The central Mexican Crassulaceae allbelong to a single evolutionary line (van Ham 1994). The

~ taxa are relatively young and show little evolutionary

0

Tylecodon paniculatus, a member of theCrassulaceae which is removed from the wild insouthern Africa because it causes stock poisoning.

divergence, which may explain why they can still behybridised forming a single huge comparium (a group ofspecies which have the potency to hybridise) of more than200 species (Uhl 1992). Many of the MexicanCrassulaceae are very local and known only from the typecollection or a few locations. Northwards along the SierraMadre the number of species gradually diminishes. The

21

adjacent regions of Baja California and the USA states ofCalifornia and Oregon constitute a secondary Americancentre of diversity independent of the central Mexicanlineage. The region comprises another 80 endemic speciesmost of which belong to Dudleya and Sedum. MostMexican and western American Crassulaceae have large,rosulate or semirosulate, very succulent, thick leaves andare often shrubby. In the USA a few annual Sedoideaeoccur, which are usually separated from Sedum, i.e.Diamorpha (1 sp.) and Parvisedum (4 “pp.). Some speciesof Echeveria and a few species of Sedum and Villadiaoccur in South America as far as north-westernArgentina.

The third centre of diversity is the Eurasian-Mediterranean region with about 220 endemic species inseven major genera (Aeonium, Aichryson, Monanthes,Rosularia, Sedum, Sempervivum, Umbilicus). A generalfeature of Eurasian Crassulaceae is the large number oftaxa with polymerous flowers, most notably Berger’ssubfamily Sempervivoideae and some groups of Sedum.The genera Aeonium, Aichryson, and Monanthes areconfined to Macaronesia, except for four species ofAeonium occurring in north-west Africa, eastern centralAfrica, and on the Cape Verde Islands. The CanaryIslands are notable for the high number of endemicspecies of these three genera, with over 60 spp. in total.Sempewivum (c. 40 spp.) is confined to the montane andalpine zones of the mountains of southern Europe andthe Near East. All species of Sempewivum are rosulateand much sought after as ornamentals. RosuZaria agreeswith Sempewivum in habit and ecological preference. It ismainly confined to the Near East. The Eurasian-Mediterranean Sedum flora is further characterised by alarge number of annual or hapaxanth species (over 40percent). Umbilicus is notable for its distinctlypedunculate, peltate leaves, racemose inflorescences, andsympetalous flowers. Umbilicus, Pistorinia (2-3 spp. with atubular corolla), and a few sympetalous species nowincluded in Sedum, used to be classified in Berger’sotherwise s t r i c t l y S o u t h A f r i c a n s u b f a m i l yCotyledonoideae.

The fourth centre is located in eastern Asia andcomprises about 200 endemic species in five major genera(Hylotelephium, Orostachys, Rhodiola, Rosularia, Sedum).A remarkable feature of the Asian Crassulaceae is thepredominance of taxa with annual, leafy, flowering shootsin combination with a rhizome (Rhodiola, c. 40 spp.,Sedum sect. Aizoon, > 15 spp.) or tuberous roots(Hylotelephium, c. 30 “pp.). Rhodiola is mainly found athigh altitudes in the Himalayas, whereas Hyloptelephiumand Sedum sect. Aizoon occur at less elevated and moremesic habitats from south-western China to eastern Asia.Many species of Rhodiola are dioecious, though theirsexual differentiation is apparently not based onchromosomal differences (sex chromosomes).

Crassulaceae generally occupy arid and semi-arid,usually rocky habitats. Only very few taxa are, secondarily,

adapted to a mesic or aquatic environments, the moststriking example of which are the semi-aquatic species ofTillaea. Although Crassulaceae are generally well-adaptedto survive extended periods of drought, they are, rathersurprisingly, largely absent from true desert regions.Throughout their area they are most abundant in sub-montane and montane regions, although they can befound from sea-level to over 4000 m in the tropics (Se&1112ruwenzoriense in East Africa). Apart from theirhorticultural value, Crassulaceae are of little or noeconomic importance. The succulent leaves aresometimes prescribed for healing wounds in folk medicineand the rhizomes of Rhodioln rosea have some use in folkmedicine. A few species have found odd applications, forexample, decoctions of Aeonium glutinosum have beenused to reinforce fishing nets on Madeira. NowadaysCrassulaceae are not used for food, though S e d u mrupestre was recommended for salad in medieval herbals.The juicy appearance of the leafy parts of manyCrassulaceae may appear rather attractive in aridenvironments, however, the succulent leaves arecompletely tasteless or bitter-tasting and are generallyeven avoided by cattle. “Not even the goats eat it” as anAnatolian peasant once told me pointing to S e d u mpallidurn. The species which contain alkaloids orbufadienolides, are generally unpalatable or evenpoisonous.

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Threats

Most species of Crassulaceae occur in rocky places whichare not prone to the threats of habitat destruction.Reforestation of montane habitats may locally reducetheir area, but is in general of little consequence.Development of mass-tourism in fragile alpine habitats oron small oceanic islands is a more acute threat and couldseriously imperil the narrow endemics frequentlyoccurring in these places. For example, of the threeaccessible populations of Sedum fusiforme on Madeira,two sites have fallen prey to extension of touristaccommodations. The few Crassulaceae occurring inarable places are all very much endangered. In addition tothe aforementioned Madagascan BryophyZZum species,some rare annual Sedoideae occurring in seasonally wethabitats could be endangered by increased cultivation, forexample, Sedum assyriacum of the Near East.

EuphorbiaceaeSusan Carter-Holmes

The Euphorbia Family (Euphorbiaceae Juss.) is a verylarge family, the sixth largest among flowering plants,consisting of over 300 genera and 5000 species in severaltribes. It is distributed throughout the temperate andtropical regions, but with the greatest representation inthe humid tropics and subtropics of both hemispheres. Inhabit the family is very diverse, with plants ranging fromprostrate, erect, or scandent annual or perennial herbs, toshrubs and trees, with leaves compound or simple, largeor minute. It is nevertheless characterised by small,unisexual flowers, and a superior, usually 3-celleddehiscent fruit with each cell containing 1 or 2 seeds.

The tribe Euphorbieae contains genera with manysucculent-stemmed species, some of these also withsucculent leaves, native in the tropics and subtropics ofthe Old World, with a few from the New World. The onlygenus outside the tribe with species that could be calledsucculent is Jatropha, a large genus confined almostexclusively to the tropics. A few species, primarily fromCentral America, have been introduced into cultivationand have become naturalised in some parts of the OldWorld. A few are caudiciform, possessing enlarged fleshystems and usually brilliant red flowers, the most widelycultivated of which isJ podagrica. A limited interest in themore woody, xerophytic species from the semi-desertareas of east tropical Africa, including Somalia, wherethey form a major part of the flora, has currently beenaroused among hobbyists, after several recentintroductions and the publication of treatments byRadcliffe-Smith (1987a, 1987b). However, these forms areusually difficult to propagate except by seed and areunlikely to become popular.


The Euphorbieae is a well-defined tribe, including thevery large genus Euphorbia, with a world-wide distributionin temperate and tropical zones; and 10 other smallgenera confined to tropical and subtropical regions. It ischaracterised by a specialised inflorescence termed a‘cyathium’, consisting of a cup-like involucre enclosingnumerous male flowers surrounding a solitary femaleflower; the male flowers are reduced to single stamens,and the female flower to an ovary with the perianthreduced to a rim, occasionally lobed. The cyathia arefurther arranged in usually dichotomous cymes. Milkylatex, which is often extremely caustic, is always present.Several genera are composed entirely of succulentspecies, namely Pedilanthus, Synadenium, Molzadenium,and Elaeophorbia, all of which are of interest to thehobbyist grower.

PediZanthus is a genus of shrubs and small trees withleafy, fleshy branches and extremely zygomorphic, beak-like cyathia usually coloured red. A thorough treatmentby Dressler (1957) includes 14 species centred in the drierregions of Central America, and extending into thenorthern part of South America and the West Indies. Onespecies, P. tithymaloides has been introduced as anornamental, and has become naturalised in many parts ofthe Old World. This and P. macrocarpa from BajaCalifornia are the most widely cultivated.

The genus Synadenium, characterised by an entire,spreading glandular rim on the involucre, is confined indistribution to east and southern tropical Africa consistingof about 20 closely related species, difficult to identify, themajority of which are centred in Kenya and Tanzania. Anaccount of the East African species has been given byCarter (1988a) for the Flora of Tropical East Africa, andLeach worked on the southern species for FloraZambesiaca. All are shrubs or small trees with semi-succulent branches and large fleshy leaves. Theirtaxonomy is difficult to work out, differences betweenthem relying primarily on leaf and cyme-branchingcharacters. Branch-cuttings are readily rooted, and red-leaved forms are commonly cultivated as ornamentals intropical regions world-wide, as well as less popularly bythe hobbyist. A few species appear to have limiteddistributions, but most are fairly widespread, usually indeciduous woodland, and are threatened only byincreasing human population pressures.

Monadenium contains over 50 species, distributedthroughout the eastern tropical regions of Africa, fromnorthern Somalia southwards to the Transvaal, themajority endemic in Kenya and Tanzania. All are limitedin distribution, occurring in dry grassland and opendeciduous woodland or scrub, as small, widely scatteredor sometimes solitary populations. A monographicaccount was given by Bally (1961) with the east Africanand Somali species later revised by Carter (1988b,c, 1993).Leach worked on the fewer southern species for FloraZambesiaca. The genus is distinguished by an entire

23

horseshoe-shaped involucral gland with a deep rimextended to protect the ovary which is exserted throughthe notch. Persistent bracts envelope the involucre andare sometimes large and showy. Habit ranges from smallgeophytes with large fleshy roots, to herbs with thick,succulent stems covered in tubercles, and small trees withoften spiny stems and large fleshy leaves. All species areattractive to the hobbyist. Most are easily propagatedvegetatively, and have been in cultivation for many years,often from the original introductions made by Ballyduring research for his monograph. Several species weredescribed from single specimens and are insufficientlyknown, especially the small, often insignificant geophytesfrom the Brachystegia woodlands of southern Tanzania,Zambia, and Malawi. These are not threatened in thewild, but the thickly succulent stemmed species from drierareas farther north are increasingly vulnerable fromhabitat destruction by a growing population, or byovergrazing.

By far the largest genus is Euphorbia, with nearly 2000species world-wide. Because of the unifying structure ofthe cyathium, classification of this large and otherwisediverse aggregate of species has always been difficult andno system proposed so far has proved entirely satisfactoryon a world-wide basis. About 650 species are trulysucculent, confined to the tropics and subtropics, and withrelatively few exceptions, occur in the drier regions of theOld World, well adapted to survive in often extremehabitat conditions. Their growth habit is very diverse,from small herbs to shrubs and large trees. Many morespecies are semi-succulent, with somewhat fleshy stems,or fleshy leaves, or with a more-or-less fleshy or tuberousrootstock. These are being increasingly adopted by thehobbyists as ‘succulents’ worthy of culture, but if theywere included in any overall treatment of succulentmembers, then all related species, covering the majority inthe genus, would have to be taken into account. Since allsucculent Euphorbia species are listed under Appendix IIof CITES, with a few geophytes from Madagascar onAppendix I, this borderline leads to problems in decidingwhich species should be included as succulent in the truesense.

Distribution

Succulent species in the New World, specifically Centraland South America, are relatively few, and not related tothose of the Old World. Several species, with erect, more-or-less cylindrical stems, small leaves and often largecyathial bracts, are mostly long established in cultivation,such as Euphorbia pteroneura from Mexico, and E. sipolisiiand E. phosphorea from Brazil. Several distinct speciesrelated to the latter have recently been discovered inBrazil but are not yet generally known in cultivation. E.misera from Baja California, Mexico, is a woodyxerophyte which has nevertheless long been accepted bythe hobbyists as a ‘succulent’.

There are a number of tree species with paired spines,

and a small group of geophytes related to E. jksiformis,from India and Southeast Asia that are finding increasingpopularity. The tree species are not well knowntaxonomically, but work is currently being done on themfor the Botanical Survey of India. The geophytes, withseveral probably distinct taxa, each have a limited rangeexcept for E. fusiformis itself, but are reported to bereasonably common. So far, there seems to be very littleartificial propagation of these, with most plants incirculation among hobbyists in India and elsewhere beingof wild origin. This is a group on the borderline ofsucculence, the only really succulent part of the plantsbeing their semi-woody or somewhat fleshy root. They areusually listed as caudiciform.

Madagascar has its own endemic group of species inthe subgenus Lacanthis, typified by stipules flanking theusually large leaf-scars, and modified as spines which areoften much divided into bristly fringes. They range inhabit from fleshy, thick-stemmed, shrubby herbs togeophytes. All are of interest to the hobbyist, andnumerous forms have recently been described as distinctspecies by Rauh in the journals of the German andAmerican cactus and succulent societies. Almost all arerestricted in distribution, mostly in the dry deciduouswoodland of the southern part of the island. They haveapparently been declining rapidly in numbers fromhabitat destruction, as well as being collected for export.There is no excuse for this, as they all seem easy topropagate, especially from seed, and several have beenwell established in cultivation for many years. The mostendangered are the geophytes, which are now listed onAppendix I of CITES.

Another group are the ‘pencil’ euphorbias, withsucculent, cylindrical, leafless branches related toEuphorbia tirucalli, and all trees or shrubs. The onlyspecies of any significant interest to the hobbyist is E.stenoclada, in cultivation for many years. Species areendangered only from habitat destruction. Rauh has givena good account of all the Madagascan succulent species invarious issues of The Euphorbia Journal (1983-1993).

One species related to this last pencil group is E.sarcostemmoides, which is reported to be a Rare endemicof the drier regions of Australia, with a scattereddistribution, and is sometimes found in hobbyistcollections. There are no other succulent species inAustralia, but E. tannensis, a fairly common, weedy herb,is beginning to find popularity by virtue of its tuberousroot. It has been adopted by hobbyists as a succulent, butshould not legitimately be included.

On the African continent there are three main typesof succulent Euphorbia: the tuberculate stemmed species,the pencil stemmed species, and the pair-spined species.The tuberculate-stemmed species are confined almostentirely to southern Africa in Cape Province, with a highincidence of endemism. They range in form from tall,leafy stems to dwarf and much-branched plants formingdomed cushions, to solitary or sparsely branched

24

individuals with very thick stems sometimes reduced to aglobose body. The inflorescences are persistent on manyspecies with the peduncles hardening into woody spines.In this group are some of the first succulents to beintroduced into cultivation, including E. obesa, E.meloformis, and E. bupleurifolia, all represented in almostevery nursery and hobbyist collection but limited indistribution and endangered in the wild. Overcollectionhas been, and possibly still is, a factor here, but habitatdestruction due to farming, mining operations, and road-building plays a major role. In many instances it is notpossible to determine conservation status, as habitats areknown to be on land where entry is restricted and in somecases forbidden. This is especially true of the mineral-richareas to the west into Namibia. In this area many of thecylindrical, pencil stemmed, leafless, spineless speciesoccur, usually as shrubs, the smaller ones, often endemic,finding some favour in the horticultural trade. They areprobably not vulnerable except perhaps from habitatdestruction, as propagation from cuttings is generallyeasy. There are also numerous pair-spined species in thisregion, mostly large, slow-growing shrubs, endemic in thisextremely arid zone which extends northwards intoAngola.

For all species of this southern African region, White,Dyer and Sloane’s two volumes on the succulentEuphorbieae (1941) is still the major work. More recently,Leach produced a number of papers describing manymore species, especially from the western areas, includingAngola. Several articles have also appeared in TheEuphorbia Journal, with valuable information ondistribution and conservation status (1981-1993).

Farther north, into tropical Africa and the ArabianPeninsula, there are some species in the pencil group, allof which are common and widespread, especially in thedrier regions. Euphorbia tirucalli itself is the mostcommon endemic in central tropical Africa butintroduced and naturalised throughout the tropics world-wide. It is used extensively for hedging purposes sincecuttings root readily and quickly grow to form animpenetrable hedge. It is not easily eradicated and insome places has spread to the detriment of the naturalvegetation, not only in Africa but also elsewhere intropical regions. As a succulent species it is listed underAppendix II of CITES and thus protected, but can alwaysbe easily distinguished from ‘look-alike’ species by itsdistinctively striated branches.

However, the pair-spined succulents provide thegreatest number of species north of South Africa. Therange in habit is very wide from trees and shrubs, tosucculent-stemmed herbs. They are found in abundancethroughout the drier regions, from Angola eastwards toMozambique, and northwards through East Africa toEthiopia and Somalia, with a few peripheralrepresentatives in the Arabian Peninsula and West Africa,including the Canary Islands, as well as in southern Africaand those tree species already mentioned in India and

Asia. Trees are generally less specialised for survival in anarid environment, and as such have the widestdistribution, being found from just above sea level to itshighest altitudes of over 3000 m (E. ampZiphyZZa). A genusof tree species, with large, fleshy leaves, Elaeophorbia,should be mentioned here, distinguished from Euphorbiaby its indehiscent drupaceous fruit, which has a thick,persistently fleshy exocarp and no sign of a perianth belowthe ovary. Two distinct species occur along forest edges,from West Africa eastwards to Uganda. Many authorsconsider the genus to be synonymous with Euphorbia.

Shrubs of this pair-spined group occur mostly indeciduous bushland, with fewer, usually leafy species inthe more humid regions of West Africa, but in semi-desert areas characterised by the development of anextremely robust spinescence. The majority of the smaller,succulent-stemmed herbs, often with a large, fleshy ortuberous root, occur mostly in exposed situations, usuallyamong rocks, and colonising the most extreme habitatconditions. It is this group which contains the greatestnumber of species and the greatest number of endemicswith extremely limited distributions. Many of thesespecies occur in East Africa, particularly in centralTanzania and the Rift Valley region of Kenya, but thegreatest concentration is undoubtedly in the Horn ofAfrica in Somalia and the Ogaden region of easternEthiopia.

Threats

Throughout these tropical regions, the greatest danger tothe survival of all these species lies in habitat destructionfor spreading agriculture, for charcoal burning, or inSomalia especially, by overgrazing. Overcollecting isapparently not, and is unlikely to be because of localgovernment restrictions, a hazard in any of these areas.

The remainder of the succulent species from areasnorth of southern Africa, number a very few non-spiny,caudiciform members related to E u p h o r b i aZongituberculosa, or to the southern African species E.trichadenia, occurring in sparse, open grasslands, and allof which are widespread and unlikely to be endangered.Apart from these, most of the fleshy-stemmed herbaceousor sometimes shrubby, leafy species from the CanaryIslands, related to E. atropurpurea, E. mellijka, and E.balsamij’era are generally considered as succulents.Several of these are endemic in the Canaries, andendangered, so warranting protection, but whether theyshould be regarded as succulent species, when other morecommon but closely related species are not, is open toquestion.

The most recent account for west African species isgiven by Keay in the Flora of West Tropical Africa (192s)and accounts of the East African and Somali species aregiven by Carter in the Flora of East Tropical Africa(1988) and the Flora of Somalia (1993). M. G. Gilbert’saccount for the Ethiopian species will appear in Volume 4of the Flora; and L. C. Leach worked on the species

25

covered by Flora Zambesiaca. The Arabian species willappear in the Flora of Arabia, edited in Edinburgh; andan account of the Canary Island species, with theirconservation status, is given in D. and Z. Bramwell’spocket flora (1974) and the recently published Red List ofthe Canary Islands (Gomez-Campo 1997).

PortulacaceaeMaike Gerbaulet

The Portulacaceae Juss. is a medium-sized familycomprising annual or perennial herbs and subshrubs tovery large shrubs. Most genera are more or less leaf-succulent, but some also have succulent stems(Portulacaria, &aria). The main characteristic feature ofthe family is the 2-phyllous floral involucre (sometimesconsidered as sepals). Some genera show scales or hairs atthe leaf bases (regarded as stipules by some authors),namely Anacampseros, Grahamia, Portulaca, Talinaria,Talinopsis, Talinum, and Xenia. The mostly bisexualflowers are usually ephemeral and self-fertile, but somespecies are dioecious (e.g. Ceraria spp.). The ovary ismostly superior, but semi-inferior in Portulaca.

The major centres of distribution are North and SouthAmerica, Africa, and Australia. In addition, some speciesoccur in areas of tropical Asia with well establishedAfrican biogeographic affinities, or in arctic Asia. Onegenus iserestricted to New Zealand and Kerguelen Island.Furthermore, there are several cosmopolitan weeds.

Several genera or species are cultivated asornamentals, Anacampseros spp. and Portulacaria afia areoften found in succulent collections. Claytonia virginica(spring beauty), some Calandrinia s.1. spp., and Portulacagrandiflora are common garden ornamentals. Lewisia spp.are grown in rock gardens. Some species serve as food orin the preparation of medicine, the roots of someAnacampseros spp. are used to brew beer; the leaves ofsome species are sometimes enjoyed as a snack. Thestarchy roots of Lewisia rediviva (bitter root) are eaten asa vegetable. Montia j’ontana, M. perf’oliata (winterpurslane), Portulaca oleracea ssp. sativa (purslane),Talinum fruticosum, and T. triangulare (with a purslane-like taste) are used as potherbs. Portulaca oleracea islocally also used as a medicinal plant. Portulacaria afia iscultivated as a fodder plant in South Africa.


Several treatments of single portulacaceous genera orparts thereof have been published during the pastdecades, McNeil1 (1975) revised the Montia and Claytoniageneric complex. The more recent account on Montia byLourteig (1991) focuses on the species of the southernhemisphere. Lewisia has been revised by Mathew (1989).Several authors worked on Portulaca; the treatment byLegrand (1962) covered mainly the American species withmany new names published. Geesink (1969) studied the

Indo-australian and Pacific species. The latestinvestigation was by Nyananyo (1987) who again confinedhimself to selected species. A group of portulacasrestricted to Australia, Africa, and some areas in betweenis currently under investigation by Gilbert and West.Several papers have been published concerning the statusof single species. The infrageneric classification as well asthe delimitation of species remain unresolved.

Concerning the systematic position of the endemicAfrican and Madagascan genera Calyptrotheca, k-aria,Portulacaria, and Talinella and the small genera Hectorellaand Lyallia (sometimes segregated under the nameHectorellaceae) several papers have been published byNyananyo (1986a, b, c, 1989, 1990). As a result, both lattergenera were united and (re)included in the Portulacaceae.The remaining genera were suggested to belong to thePortulacaceae rather than to the Basellaceae.Nevertheless, their position within the Portulacaceaeremains unresolved.

The southern African species of Talinum have beenrevised by Tolken (1969). However, a revision of thewhole genus, or even the American species, is wanting.Besides, the new genus Amphipetalum recently describedfrom Argentina (Bacigalupo 1988) seems to be closelyrelated to Talinum. Also, the relationship betweenCalyptrotheca and Talinum should be re-evaluated.

In recent years, there have been several attempts tore-examine the generic relationship within the family. Asynoptical classification of Portulacaceae was proposed byMcNeil1 (1974). This showed, however, that manyquestions concerning the systematics of portulacaceousgenera still remain to be answered. The starting point forthorough changes in the classification and circumscriptionof genera within the Portulacaceae was the cladisticanalysis of Carolin (1987). One of the most importantresults of his analysis was that the large and most diverseportulacaceous genus Calandrinia s.l., is unnatural.Hershkovitz (major publications concerning thetaxonomy, 1991, 1992, 1993) revised the whole genericcomplex and assigned the species to five genera,Schreiteria, Rumicastrum, Calandrinia s.str., Cistanthe, andMontiopsis. Furthermore, Carolin’s cladogram served as abasis for a revision of the genus Anacampseros and relatedgenera including a thorough evaluation of their systematicposition within the Portulacaceae (Gerbaulet 1992a). As acorollary, the monotypic Argentinian genus Xenia wassegregated from Anacampseros.

At present, most of the generic relationships withinthe Por tu lacaceae a re unreso lved . Also thecircumscription of the family must be reconsidered.Various authors pointed out that the Portulacaceae, aspresently understood, are in all probability paraphyleticand that. the Basellaceae, Didiereaceae, Hectorellaceae,and presumably even Cactaceae have to be considered toresolve the phylogenetic relationship within the first.

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Generic distribution known about the conservation status of the East African

The main distribution areas of the Portulacaceae areNorth and South America, Africa, and Australia,although some genera are also found elsewhere. Theseareas can be sorted into two groups: the first mainlycomprises areas in North and South America east of thecordilleras, in eastern and southern Africa, and/or inAustralia. The second group mainly includes areas inNorth and South America west of the cordilleras.

The major members of the first group areAnacampseros, Portulaca, and Talinum, the latter beingrestricted to America and Australia. Portulaca, showing amore tropical distribution, is also found in those areas oft rop ica l As ia wi th we l l e s tab l i shed Afr icanbiogeographical affinities, like tropical Arabia, southernIndia, and Sri Lanka. Furthermore, some weedy species ofPortulaca and Talinum are cosmopolitan (e.g. P. oleracea,T. paniculatum). Some smaller genera are confined tocertain parts of the first set of distribution areas:Talinaria and Talinopsis are endemic to Mexico.Amphipetalum, Grahamia, Schreiteria, and Xenia occur ineastern South America. Calyptrotheca (eastern Africa),Ceraria, Portulacaria (both southern Africa), and Talinella(Madagascar) are native to Africa including Madagascar.Rumicastrum is endemic to Australia.

Calandrinia s.str. (including Monocosmia), Cistanthe(sensu Hershkovitz 1992, including Calyptridium, Lewisiatweedyi, Philippiamra, and Sparguea), and Montia are themain genera of the second distribution block. SomeMontia species are cosmopolitan weeds (e.g. M. fontana).Claytonia and Lewisia are native to western NorthAmerica including Alaska. They share an arctic totemperate distribution with Claytonia extending intoarctic Asia. Lenzia and Montiopsis occur in western SouthAmerica.

Lyallia (including HectoreZZa) is endemic to NewZealand and Kerguelen Island. At present, no agreementhas been reached whether to sort these to the first or thesecond set of distribution areas.

Threats and conservation status

Of all the Portulacaceae, Anacampseros spp., Cistanthe(formerly Lewisia) tweedyi, Lewisia cotyledon, L. maguirei,and L. serrata are protected by the CITES Appendix IIlisting.

Regarding the genera endemic to or showing a majordistribution centre in Africa and Madagascar, only little is

Calyptrotheca and the Madagascan Talinella. T h eendangered or threatened southern African taxa are listedin the Red Data List of’ southern A.frican plants (Hilton-Taylor 1996b).

In Africa, Portulaca and Talinum consist mainly ofwidespread species. Some species of both genera arcprobably not endemic to Africa but became naturalisedafter being introduced as garden ornamentals or weeds.Talinum species are not threatened in Africa. Portulacaappears to comprise a few very widespread speciescomplexes which are often segregated into several tomany minor species. Some of these microspecies arcrecorded to be threatened (e.g. Portulaca trianthemoidesin South Africa), but the species complexes as a whole arcnot.

The genera Ceraria and Portulucaria consist only of afew species each. Although ‘real succulents’, most of themhave not been very popular among horticulturists untilrecently. An exception is Portulacaria a@a which has beenin cultivation for decades. Most of the species have only arather limited distribution area. This is especially true forPortulacaria armiana which is known from a couple oflocalities only. Since this recently described species seemsto have become the fashion among collectors, it should beclassified as Vulnerable. The other species, however, arefairly common and not threatened.

Conversely, all African Anacampseros species (22currently recognised) have tremendous horticulturalvalue. Although single species are fairly widespread, manyspecies show a restricted distribution. Generally, the morewidespread species occur only sporadically, whereas therestricted ones may be locally abundant. Since the maindistribution area of Anacampseros is not subject to large-scale agricultural activities, the major threat is imposed bycollectors. In the eastern parts of South Africa, however,agriculture does have an impact on the conservationstatus of Anacampseros. In this region, A~%a~~anzilsei-~~sspecies are nowadays more or less restricted to rockyoutcrops or mountainous habitats, but herbarium recordsshow that they have been far more widespread in therecent past. In northern Namibia, overgrazing by cattleleads to the spreading of thornbush savanna (Walter1964; Walter and Breckle 1984) which in turn threatensthe endemic Anacampseros species (Gerbaulet 1992b). Onthe whole, most species should be classified as Rare orVulnerable, some even as Endangered (14 in total), whilethe remaining eight do not seem to be threatened.

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Chapter 2

Conservation Measures

National legislationSara Oldfield

In general succulent plants are relatively distinctive withinnational floras, and due to their horticultural attraction,wild populations have been widely sought after andexploited. As a consequence, where the laws relate tocollection and trade, succulents may be quite wellrepresented in national conservation legislation for plants.Nevertheless, the lack of basic inventory and assessmentof conservation status for succulent plant species limitsthe development of species conservation legislation inmany countries. This is particularly the case in SouthAmerica where, in general, laws protecting plant speciesrestrict collection and export of a wide range of plants butrarely protect individual succulent plant species.Madagascar is another important succulent-rich country

which has no specific legislation protecting rare andthreatened succulent species. The USA currently has themost comprehensive legal protection for succulent plantspecies in the wild. The Endangered Species Act (1973)(ESA) currently protects 24 native cacti and two succulenttaxa, listed as Threatened or Endangered as shown inTable 2.1. The ESA prohibits interstate or internationaltrade in listed plants without a permit. The removal oflisted plants from lands under federal jurisdiction is alsoprohibited without a permit.

Habitat conservation is covered by section 7 of theESA. Federal agencies are required to ensure that anyactions they fund, authorise, or undertake are not likely tojeopardise listed species. Another legal requirementunder ESA is the preparation of recovery plans forindividual species. These include positive measures toenhance the survival of listed species in the wild throughmanagement of wild populations and reintroduction

Table 2.1 Threatened succulents and cacti listed on the US Endangered Species Act (as of30 September 1996)

Taxon

AgavaceaeAgave arizonica

Common name Distribution

Arizona agave U.S.A. (AZ)

Listing’

E

CactaceaeAncistrocactus (= Echinocactus Tobusch fishhook cactus U.S.A. (TX) E

= Mammillaria) tobuschiiCereus eriophorus var. fragrans Fragrant prickly-apple U.S.A. (FL) EAstrophytum (= Echinocactus} Star cactus U S.A. (TX) j Mexico E

as teriasCoryphantha (= Cochiseia = Cochise pincushion cactus U.S.A. (AZ), Mexico (Sonora) TEscobaria) robbinsorum

Coryphantha (=Escobaria = Nellie tory cactus U.S.A. (TX) EMammillaria) minima(= C. nellieae)

Coryphan tha ramillosa Bunched tory cactus U.S.A. (TX), Mexico (Coahuila) TCoryphantha scheeri var. Pima pineapple cactus U.S.A. (AZ), Mexico (Sonora) E

robus tispinaCoryphantha sneedii Lee pincushion cactus U.S.A. (NM) T

(= Escobaria = Mammillaria) var.leei

Coryphantha (= Escobaria = Sneed pincushion cactus U.S.A. (NM, TX) EMammillaria) sneedii var. sneedii

Echinocac tus horizon thalonius Nichol’s Turk’s head cactus U.S.A. (AZ) Evar. nicholii

Echinocereus chisoensis Chisos Mountain hedgehog U.S.A. (TX) T(=reichenbachii) var. chisoensis cactus

Echinocereus fendleri Kuenzler hedgehog cactus U.S.A. (NM) E(=hempe/ii of authors, not Fobe)var. kuenzleri

Echinocereus lloydii Lloyd’s hedgehog cactus U.S.A. (NM, TX) E(= E. roetteri var. Iloydii)

28

Echinocereus reichenbachii var.albertii (= E. melanocen trus)

Black lace cactus U.S.A. (TX) E

Echinocereus triglochidiatus var. Arizona hedgehog cactus U.S.A. (AZ)arizonicus (= E. arizonicus)

Echinocereus viridiflorus var. Davis’ green pitaya U.S.A. (TX)davisii (= E. davisii)

Echinomastus (= Echinocactus, Lloyd’s Mariposa cactus U.S.A. (TX), Mexico (Coahuila)= Sclerocac tus, = Neolloydia)mariposensis

Harrisia (= Cereus) portoricensis Higo chumbo U.S.A. (PR)Leptocereus grantianus None U.S.A. (PR)Opun tia treleasei Bakersfield cactus U.S.A. (CA)Pediocactus (= Echinocactus, Siler pincushion cactus U.S.A. (AZ, UT)

= Utahia) sileriPediocactus (= Echinocactus, Peebles Navajo cactus U.S.A. (AZ)

= Navajoa , = Toumeya, = Utahia)peeblesianus var. peeblesianus

Pediocactus (= Tourneya) bradyi Brady pincushion cactus U.S.A. (AZ)Pediocactus (= Tourneya) Knowlton cactus U.S.A. (CO, NM)knowltonii (= P. bradyi var.knowltonii)

Pediocactus despainii San Rafael cactus U.S.A. (UT)Pilosocereus (= Cereus) robinii Key tree-cactus U.S.A. (FL), CubaSclerocactus (= Coloradoa, = Mesa Verde cactus U.S.A. (CO, NM)Echinocactus, = Pediocactus)mesae-verdae

Sclerocactus (= Echinocactus, Uinta Basin hookless cactus U.S.A. (CO, UT)= Pediocactus) glaucus (= whippleivar. glaucus, =subglaucus,= franklinii )

Sclerocactus (= Pediocactus) Wright fishhook cactus U.S.A. (UT)wrightiae

Dudleya se tchellii Santa Clara Valley dudleya U.S.A. (CA)Dudleya traskiae Santa Barbara Island liveforever U.S.A. (CA)Sedum in tegrifolium ss p . leedyi Leedy’s roseroot U.S.A. (MN, NY)

1 E =Endangered: any species in danger of extinction throughout all or a significant portion of its range.T =Threatened: any species likely to become an endangered species within the foreseeable future throughout all or

a significant portion of its range.

E

E

T

TEET

E

EE

EET

T

E

EET

schemes, recovery meaning restoration of the populationto a point where it is a “viable self-sustaining componantof their ecosystem so as to allow downlisting” (US Fishand Wildlife Service 1990). A 1988 amendment to theESA requires that recovery plans for listed species bedeveloped and implemented by the Department of theInterior. Reports are made to Congress every two years.These recovery plans are intended to provide a blueprintfor private, Federal and State interagency cooperation onimplementation.

At present the ESA is under some threat. In 1995 amoratorium on new listing was called, but a year latercertain cases were pushed through. Several bills havebeen introduced to Congress that aim to weaken theprotection of national endangered plants; these will makeit more difficult for the USA to fulfil its obligations withCITES (De Ferrari 1996).

Another Federal law in the USA, the Lacey Act, givesFederal support to state conservation laws. Since 1981,the Lacey Act has prohibited interstate trade or export ofwild native plants collected or possessed in contraventionof the State or, in the case of Indian lands, the

Reservation, of origin. States which have legislationregulating the collection of cacti and succulents areArizona, California, New Mexico, Nevada, and Texas.

Zimbabwe is another country which has good legalprovisions protecting succulent plants both fromcollection pressures and habitat destruction. The Parksand Wildlife Act, 1975, as amended lists the genus Aloeand ten other succulent species as specially protected. TheAct also allows for the constitution of botanical reservesto protect rare or endangered indigenous plants orrepresentative plant communities growing naturally in the

.* *wild.

Unfortunately, protective legislation for succulentplants is not enforced in Zimbabwe due to lack ofpersonnel and the absence of any active and full-timeinspectorate. No reserves have yet been createdspecifically to protect representative communities ofsucculent plants which are specially protected by law. Arecommendation to create a network of succulentreserves under the Parks and Wildlife Act, 1975, is givenin Chapter 4 of this Plan, the Action Proposals.

In South Africa, the Forest Act 122 of 1984 is a

29

national statute which gives protection to a number oflisted succulents. The most important legislation for theprotection of native plant species is, however, provided bythe four provincial ordinances (Glavovic 1993). Underthese ordinances, plants on or along public roads areprotected and no native plant can be picked without thelandowner’s permission. Possession and trade inindigenous plants is generally regulated and certainspecies are given special protection.

In Natal all native flowering plants are regulated withcertain species classified as specially protected. TheTransvaal also has a list of specially protected species,together with lists of endangered and rare species.Importation and exportation of endangered and rarespecies is controlled. The Cape legislation listsendangered flora and protected flora. Succulent speciesgiven special protection under the South African stateordinances are listed in Table 3.8 in the Southern AfricaRegional Account in Chapter 3.

A recent review of plant conservation legislation inSouth Africa has pointed out that the existing laws do nottake into account traditional rights and harvestingpractices. Controlled access to plants used, for example,for medicinal purposes should be part of policy toencourage sustainable utilisation. Furthermore, it ispointed out that an integrated national legal system forflora conservation is needed with consistency throughoutthe regions. Perhaps of most importance, greateremphasis should be placed on in situ conservation so as topreserve representative and important plant communitieswherever possible.

The development of conservation legislationprotecting rare and threatened succulent species remainsa priority in many countries, particularly in Madagascar.Ideally, legislation should aim to enhance theconservation status of individual species through thedevelopment of recovery programmes, as well as limitinghabitat destruction and overexploitation. Implementationand enforcement of existing legislation also need to beaddressed. One catalyst for improving national legalprotection for succulents is the development ofincreasingly comprehensive international legal obligationsto protect threatened species and their habitats.

International legislationThe Convention on Biological Diversity

The Convention on Biological Diversity (BiodiversityConvention) was signed by 153 States, together with theEuropean Community, during the UNCED meeting inJune 1992. The objectives of the Convention are theconservation of biological diversity, the sustainable use ofits components, and the fair and equitable sharing of thebenefits arising out of the utilisation of genetic resources.Parties to the Biodiversity Convention are required toidentify components of biological diversity important for

conservation and sustainable use; develop nationalstrategies, plans or programmes for the sustainable use ofbiodiversity; establish protected area systems; develop ormaintain threatened species legislation; and furtherbiodiversity conservation by various other specifiedmeans.

The Biodiversity Convention will enhance theconservation of cacti and succulents in various countriesthrough the identification of threatened species anddevelopment of the conservation measures outlinedabove. Country biodiversity studies have been producedfor a range of countries including Kenya and Uganda as apreliminary step in implementation of the Convention.These form the basis for the development of the nationalstrategies or action plans to protect biodiversity.

The Berne Convention

This Convention, applicable to the Council of Europemember states, was signed in 1979. The Conventionprovides for the conservation of wildlife and wildlifehabitats in general and for the special protection of listedspecies. Appendix I of this convention lists 490 speciallyprotected plants including the succulents listed in Table2.2, all of which occur in the Canary Islands except forEuphorbia stygiana which is endemic to the Azores.

Table 2.2 Threatened succulent speciesin Appendix I of the Berne Convention

AgavaceaeAsclepiadaceae

Crassulaceae

Euphorbiaceae

Leguminosae

Dracaena dracoCaralluma burchardiiCeropegia chrysanthaAeon&m gomeraenseAeon&m saundersiiEuphorbia handiensisEuphorbia IambiEuphorbia stygianaLotus kunkelii

The EC Habitats Directive

Council Directive 92/43/EEC of 21 May 1992 on theConservation of Natural Habitats of Wild Fauna andFlora, the European Community Habitats Directive, aimsto contribute to the protection of biodiversity through theconservation of habitats and wild fauna and flora withinEC countries. Under the Directive a network of protectedhabitats is being created through the designation ofSpecial Areas of Conservation. The network will includethe habitats of species listed in Annex II of the Directive.Annex IV (b) designates plant species in need of strictprotection and Annex V lists plant species whose taking inthe wild and exploitation may be subject to managementmeasures. Macaronesian succulent plant speciesprotected by the Habitats Directive are listed in Table 2.3.

30

Table 2.3 Succulent species in theAnnexes of the EC Habitats Directive

ANNEX IIAsclepiadaceae

Crassulaceae

Euphorbiaceae

Leguminosae

Caralluma burchardiiCeropegia chtysanthaAeonium gomeraenseAeonium saundersiiAichtyson dumosumMonanthes wildpretiiSedum brissemoretiiEuphorbia handiensisEuphorbia lambiiEuphorbia stygianaLotus kunkelii

In addition to the regionally protected species listed inthese Annexes, parties are required to “identifyendangered or threatened species of flora or fauna withinareas over which it exercises sovereignty, or sovcrcignrights or jurisdiction, and accord protected status to suchspecies”.

The SPAW Protocol also sets out a cooperativeprogramme for protected areas in the Caribbean region.This will support the selection, establishment, planning,management, and conservation of protected areas andbuffer zones, where necessary, and the creation of aprotected areas network.

ANNEX IV (b) Controlling the tradeContains the succulent species listed above togetherwith:Agavaceae Dracaena draco

ANNEX VNo succulent species listed at present.

Sara Oldfield*

The SPAW Protocol of the Cartagena ConventionThe Protocol on Specially Protected Areas and Wildlife(SPAW) was developed under the Cartagena Conventionon the Protection of the Marine Environment of theWider Caribbean Region. Annex I of the Protocol lists 56vascular plant species which are protected from all formsof destruction or disturbance, including picking,collection, or commercial trade. As far as possible,activities which adversely affect the habitats of thesespecies will be subject to regulation. Annex III of theprotocol lists ‘harvestable’ species of flora and fauna,exploitation of which will be regulated according tomanagement plans. With respect to the selected cacti andother succulent taxa, these Annexes are far from completeand up to date (Table 2.4). Conservation of species onboth Annexes will require not only protection but alsomeasures to enable the recovery of wild populations.

Cacti and other succulent plants are of major horticulturalimportance forming the basis for a multi-million dollarinternational industry. They are sold both as generalhouseplants and as botanical specimen plants forspecialist collectors. Collection of plants from the wild forinternational trade has been one of the main threats tocertain species, and despite the development ofsophisticated propagation techniques this threat remainsa significant problem.

CITESThe principal means by which international trade insucculent plants is controlled is the Convention onInternational Trade in Endangered Species of WildFauna and Flora (CITES). This convention first cameinto force in 1976 and has now grown to become one ofthe largest and best known of all conservationconventions, with over 130 member states. The generalaim of CITES is to regulate trade in threatened wildspecies as listed in three appendices.

The Appendices

Table 2.4 Succulent species listed in theAnnexes of the SPAW Protocol

ANNEX ICactaceae

ANNEX IllAgavaceaeAsclepiadaceaeCactaceae

Echinocereus reichenbachii var.albettii

Harrisia fragransHarrisia portoricensisLeptocereus grantianusLeptocereus wrigh tiiMelocac tus guitatfiiMelocactus harlowii s. I.Pilosocereus deeringiiPilosocereus robinii

Nolina brittonianaAsclepias viridulaMelocactus intortus

Species listed on Appendix I of CITES (Annex 3) arcconsidered to be at serious risk from any commercialtrade and thus all international trade in wild plantsbetween countries who are members of the convention isbanned under the terms of the Convention. Species listedon Appendix II of the Convention are those which may bethreatened by excessive levels of trade withoutappropriate regulation. Species may also be included inAppendix II because of their similarity to morethreatened species, as an aid to enforcement.International trade in wild specimens of Appendix IIspecies is permitted under the convention but iscontrolled and monitored through a licensing system.Appendix III is used by countries which may want tocontrol trade in particular species of their own flora andfauna which are not listed on the other Appendices. It isnot currently used for any succulent plants.

*Account prepared in 1993

31

The Appendices of CITES are amended by agreementof the member states. Normally changes are madefollowing consideration of detailed proposals at theConference of the Parties which is held every two years.Changes may also be made through postal votes betweenthe meetings.

A list of the succulent plants included in theAppendices of CITES is given in Annex 3. The listing isnot fully comprehensive for horticulturally desirablesucculents or fully representative of species threatened bytrade. Some succulents were included in the originalAppendices of the Convention drawn up in 1973 beforedetailed justifications were required for each taxon listed.Subsequent amendments to the Appendices have beenbased on more thorough compilation of information, but,in general, the data available on which to base CITESproposals for succulent genera has been, and remains,incomplete.

Many succulents do not yet benefit from internationaltrade controls through CITES. These include manyrarities native to Madagascar and Africa which are soughtby collectors. There is currently a fashion within the tradefor caudiciform or swollen-stemmed plants, which occurin a wide range of families and genera. During the pastdecade caudiciform plants, such as Kedrostris,Raphionacme, Cyphostemma, Dioscorea, Adenium, andFockea, all non-CITES, together with Pachypodium and

CITES export permit, Peru.

Euphorhia caudiciforms, have become popularhouseplants. Many of these plants are slow-growing incultivation and wild-collected plants are regularly offered.Certain genera and species warrant CITES listing and arecommendation for development of proposals is given inChapter 4. Further research is necessary to determine theimpact of trade on other succulents and their suitabilityfor CITES listing.

Artificially propagated plantsCITES defines artificially propagated plants as grownfrom seeds, cuttings, or propagules under “controlledconditions”, where the stock has been established andmaintained in such a way that does not damage thesurvival of the species in the wild. The material should bemanaged in a manner designed to maintain the artificiallypropagated stock indefinitely.

Artificially propagated Appendix I species may betreated in the same way as Appendix II species under theterms of the Convention. Artificially propagatedAppendix II species are effectively treated as if they werenon-CITES by some countries, although permittingrequirements vary. It is generally considered that someminimum documentation needs to accompany suchAppendix II species in international trade to show thatthey are not taken from the wild. One option allowed byCITES is the use of phytosanitary certificates annotatedfor CITES purposes.

Millions of artificially propagated plants of both non-CITES and CITES-listed succulent species, includinglarge numbers of those on Appendix I, are tradedinternationally. This legitimate trade does not impact onwild populations of the plants concerned and should beencouraged to stem demand for wild plants. Mostnurseries trading internationally deal only withpropagated material, but this is not always the case. Thereare frequent instances of wild plants being mislabelled asartificially propagated in order to escape CITES controls.Commercial succulent plant propagation in succulent-richcountries such as Madagascar and Mexico is not yetsufficiently well developed, as discussed in the regionalsections of Chapter 3.

A resolution was passed on nursery registration at theCITES Conference of the Parties held in Fort Lauderdalein 1994. The intention of nursery registration is primarilyto facilitate propagation and trade, is widely seen as adesirable means to simplify the licensing arrangements.This would ease the burden of paperwork for nurseriesexporting only artificially propagated material.

Implementation of CITESImplementation of CITES is through the nationallegislative and administrative procedures of memberstates. The text of the Convention and subsequentresolutions and recommendations agreed by the Partiesprovide the framework for national implementation andinternational cooperation. The convention requiresParties to appoint Scientific and Management Authorities

32

Propagation at theDesert Botanical Garden.

who are responsible for carrying out the provisions of theconvention.

Unfortunately, implementation of CITES has beengenerally weak for plants, and some countries have largelyignored the requirements of the Convention for plants.Few countries have allocated the resources to determineappropriate levels of trade in wild plant species on ascientific basis prior to issuing export permits. In othersadministrative procedures have hindered legitimate tradein artificially propagated CITES plants alienating bonafide nurseries from the CITES process. The intent of theCITES convention can be lost in excessively bureaucraticlicensing procedures, especially where these are notclearly explained to importers and exporters.

Table 2.5 lists the main countries involved in theimport and export of cacti and other succulents andprovides examples o f r e p o r t e d p r o b l e m s i nimplementation of CITES controls for these plants.

Certain general problems with enforcement of theConvention for plants are being addressed, but relativelylimited attention is given to these problems comparedwith enforcement for animals. One major problem is thatof plant identification. Few enforcement agents havespecialist botanical training and even with such training itcan be very difficult to recognise species listed under theConvention or to distinguish between Appendix-I andAppendix-II listed succulents in the same genus. Agreater problem is to distinguish between artificiallypropagated and wild-collected plants of the same species.

Some identification guides have been produced tohelp enforcement. The Italian Succulent Plant Society(AIAS) has produced an identification manual forAppendix I cacti, for example, with colour photographs.The Swiss CITES Management Authority has alsoproduced a guide, and draft identification sheets forCITES-controlled plants have been produced by the USCITES Agencies. In the UK the CITES Guide to Plants inTrade has also been published (Mathew 1994).

EC CITES regulationIn EC countries, CITES controls are implemented bymeans of Council Regulation 3626/82, which came intoforce on 1 January 1984. This Regulation goes beyond thebasic requirements of CITES by imposing the necessityfor import, as well as export, permits for CITESAppendices II and III species brought into theCommunity.

As both a major consumer of wild plants forhorticulture and a major producer of artificiallypropagated CITES species for export, it is important thatthe EC has effective means to implement the Convention.Inconsistencies remain in the enforcement, with someEuropean countries operating very lax controls on theplant trade. There is a need for a coordinated CITESinspectorate within the EC.

In order to improve implementation, the EC CITESRegulation has been subject to extensive review and arevised version has been prepared. The future of therevised legislation remains unclear. It has been proposedthat a significantly increased list of plant species will besubject to trade controls if and when the new EClegislation comes into force. A new provision of thelegislation may also be to introduce a list of plants (andanimals) for which import into the EC would bemonitored only, without a specific requirement for exportdocumentation. The intention is to build up a clearerpicture of the quantities of listed species entering theCommunity to provide an early warning system ofpotential trade threats.

Monitoring the trade in CITES succulentsOne of the major successes of CITES for succulent plantshas been to compile information on the volumes of tradein Appendix II species, providing information which is notavailable from any other source. All Parties to theConvention are obliged to submit annual reports on tradein CITES-listed species to the CITES Secretariat.

33

Table 2.5

Country

Austria

Belgium

Bolivia

Brazi I

Canada

Chile

Denmark

DominicanRepublic

France

Germany

Italy

Japan

Korea

Madagascar

Mexico

Namibia

Netherlands

Peru

South Africa

UK

USA

Zimbabwe

Main countries involved in cactus and succulent trade with some specificproblems relating to control

Import/Export

Import and export

Import and Export

Export

Export

Import and Export

Export

Import and Export

Import and Export

Import and Export

Import and Export

Import and Export

Import and Export

Import and Export

Export

Export

Export

Import and Export

Export

Import and Export

Import

Import and Export

Export

Problems relating to control in trade of cacti and succulents

Export of wild Appendix I species to Italy (Jenkins 1992). Confiscation of 49 kgof Mexican cacti in August 1993.

Mexican Appendix I cacti available in trade (Jenkins 1992).

Poor reporting of CITES plant trade. Collecting of rare plants by Europeanenthusiasts.

Under-reporting of the cactus trade. Export of large quantities of seed isdamaging certain wild populations.

A major trader in Cactaceae; few details of the trade.

Wild-collected Copiapoa have been popular in European trade. Chilean plantsexported via Peru.

Concern about ‘laundering’ of cacti and succulents collected in othercountries.

Wild-collected Mexican Appendix I cacti available in trade (Jenkins 1992).Seizure of 390 wild collected Mexican cacti at Orly airport in February 1993.

Wild-collected Mexican Appendix I cacti available in trade (Jenkins 1992).

Relatively recent development of cactus hobby has led to strong demand forwild-collected rarities. Seizure of 1490, mostly wild-collected, cacti arriving fromPeru.

Import of rare field-collected cacti.

A major trader in Cactaceae; few details of the trade.

International demand for native succulents far exceeds nursery capacity topropagate the plants. Wild plants have been exported in bulk as artificiallypropagated specimens. Uncontrolled trade in non-CITES succulents.

Major source of illegally collected cacti in trade. National legislation has notbeen effective in preventing the export of rare wild plants from the country overthe past 40 years. Recent improvements in enforcement but controls at theMexico/USA border remain difficult and specialist collectors from Europe andJapan pose problems.

Export of wild plants is prohibited but European nursery catalogues indicateavailability of both CITES and non-CITES succulent plants,

Major international trading country for succulents. Huge trade in artificiallypropagated plants but Mexican Appendix I cacti available in trade as wildplants (Jenkins 1992).

Under-reporting of CITES trade in cacti and wild cacti traded as artificiallypropagated. Seizures of large consignments of wild cacti in USA and Europerecently. A shipment of 1491 plants investigated in Palermo in 1993, showedthat more than 90% wild collected although all were claimed to be artificiallypropagated.

Concern has been expressed about the export of wild ,!?uphorbia spp. Manyother genera of the rich succulent flora are in export trade but details are notknown. Export of non-CITES succulents is a cause of concern. Seizures of wild-collected CITES succulents without permits by UK Authorities in 1994.

Licensing requirements have deterred some nurseries from export market.

Delays and expense in obtaining permits has hindered legitimate export ofartificially propagated plants.

Inadequate enforcement of national legislation results in some illegal trade inrare plants.

34

However, the standard of reporting for plants is generallypoor. Many countries, for example, report only to familyor generic level for cacti and other succulent plants, andother countries do not report on the plant trade at all.Nevertheless, a significant body of data on theinternational succulent plant trade has been compiledsince the Convention came into force.

Monitoring of the trade through review of licenceapplications and analysis of annual reports can provide anearly warning system of trade on a scale likely to causeconservation problems. This happened, for example, withthe import of CITES Appendix II Madagascan succulentspecies in the mid 1980s. One species, Pachypodiumbrevicaule, was imported in the tens of thousands fromMadagascar to Germany in 1985 and 1986 for sale in

general horticultural outlets. The plants were claimed tobe artificially-propagated but subsequent investigationdemonstrated that all were wild-collected. In the sameway thousands of wild plants of the slow-growing speciesEuphorbia cap-saintmariensis, E. cylindrifolia, E. mom tii,and E. primulifolia were imported into Europe in the mid-1980s together with large numbers of Didierea andAlluaudia in the Didieriaceae. Concern about the volumesof trade lead to the transfer of certain Madagascansucculents to Appendix I of CITES in 1989 and to thetemporary imposition of stricter importer controls on allspecies by the EC.

Data from the CITES annual reports is held oncomputer at WCMC in a form which allows for variousanalyses to be carried out. Comparisons can for example

Box 2.1 CITES trade data for cacti

Cacti are the most heavily traded group of plants recorded in CITES trade statistics. The average total number of cactirecorded in annual trade is over 13 million. This is unlikely to represent the total world trade in cacti. One wholesalenursery in the Netherlands for example, produces over 18 million cacti annually, mainly for the European market In theUSA total cactus production has been estimated at up to 50 million annually, with over 20 million produced in nurseries ofVista, California alone (Fuller 1987).

The countries with the highest levels of reported average annual trade in cacti are the Netherlands (over six million),Japan (over five million), Brazil (over one million), Korea, Canada, Spain, and Dominican Republic. The high volume oftrade in plants in most of these countries is predominantly in artificially propagated cacti produced domestically and doesnot give rise to any concern. The export of cacti from Brazil is different in that both artificially propagated specimens ofindigenous and non-indigenous species are traded together with wild-collected plants.

Brazil has several major nurseries in the south of the country which are thought to deal entirely in artificially propagatedplants. One nursery exports around five million cacti annually to a wholesale firm in the Netherlands. The Brazilian trade inCactaceae is, therefore, clearly under reported. Some Brazilian cacti are relatively difficult to propagate and are stillsought after as wild specimens. Concern about levels of trade in wild-collected plants of Uebelmannia and Discocac&~~,together with some species of Melocactus, led to the transferral of these species to Appendix I of CITES in 1992. There isalso the possibility that wild-collected plants of other genera such as Notocacfus are being exported in small quantitiesfrom Brazil.

It is thought that there is substantial under-reporting of cactus exports from other South American countries, The annualaverage reported trade for Peru, for example, is only 1037 plants, 55% of which are reported to be artificially propagated.There is an internationally known cactus nursery within the country which has regularly exported wild-collected plants incontravention of CITES during the past ten years.

The average annual number of cacti reported to be exported by Mexico is around 50,000. Of these, less than one percentare reported to be propagated. As the export of wild-collected cacti from Mexico has theoretically been banned for thepast 50 years and many of the horticulturally desirable species are threatened in the wild, this scale of trade is obviouslyof considerable concern.

The most heavily traded cactus genus is Mammillaria, one of the largest genera in the Cactaceae with around 150 validspecies and many other names in horticultural use. The centre of distribution for Mammillaria is Mexico and many speciesare confined to small areas within the country. Although the genus is very commonly cultivated, it has been reported thatwild populations are exploited to fill the commercial demand for large specimens. Worrying levels of trade have beenrecorded in species such as M. dixanthocentron which is slow growing in cultivation and Vulnerable in the wild, and M.guerreronis, which is uncommon in cultivation. Newly described Mammillaria species are particularly vulnerable tocollection. Several new species are described each year in Europe, possibly as a result of illegal export of field-collectedmaterial. Publication of locality details with description of new species attracts collectors and puts the plants at risk.

Other heavily traded cacti genera include Gymnocalycium, Echinopsis, and Notocactus. Despite their horticulturalimportance these South American genera are poorly known in the wild and are in need of taxonomic fieldwork. The regionof southern Bolivia and north Argentina has the greatest diversity of small globular cacti after Mexico and they are subjectto quite a strong demand from collectors.

Source: Oldfield (I 99 1)

35

be made of the data on transactions reported by exportingand importing countries. Discrepancies highlightproblems which may be worthy of further investigation.

At the request of the Parties, a comprehensive reviewof all CITES Appendix II trade data was undertaken in1991, reviewing all trade data for the period 1983-1989.The results of this ‘Significant Plant Trade Study’ werepresented at the Eighth meeting of the Conference of theParties in 1992. Particular attention was paid to theCactaceae and a summary of the results is presented inBox 2.1. As part of the same project, trade in the genusAloe and its parts and derivatives, has been subject toreview. This concluded that the data on levels of trade inAloe parts and derivatives contained within CITESAnnual Reports currently have limited value forconservation purposes. The only significant trade in partsand derivatives from wild populations reported to theCITES Secretariat is the trade in Aloe ferox from SouthAfrica. This trade is large but appears to be sustainableand does not currently have a detrimental impact on thiswidespread species. CITES monitoring has benefits forlong-term management of A. ferox. Other countries whichexport Aloe products derived from indigenous speciesshould report the trade in detail. Importing countriesshould also record and report the trade.

The data on levels of trade in live Aloe plants recordedin CITES statistics for the period 1983-1989, show thatthe most heavily traded species are generally ‘notthreatened’ in the wild and are commonly artificiallypropagated. Relatively small-scale trade in rarer speciesmay however be a cause for concern. Collector demand isthought to focus on South African and Madagascanrarities. A number of Madagascan species are strongcandidates for Appendix I listing.

Improvement in national reporting of trade in CITES-listed succulents is a priority in increasing theeffectiveness of the Convention. A review of nationalreporting procedures for trade in CITES plants is beingcarried out by WCMC in response to a recommendationof the ‘Significant Plant Trade Study’.

Regular review of the CITES trade data for succulentsis essential in monitoring the application of theConvention to trade in these plants. A long-term goalshould be to relate trade data to biological data andinformation on nursery production for each species ofconservation concern in order to determine managementpolicies for sustainable trade. At present there are majorgaps in the available information most notably on theimpact of commercial trade on succulent populations inthe wild. Under the ongoing CITES Significant PlantTrade process, projects are now being implemented whichrelate levels of trade to the status of rare species in thewild. Recent projects carried out in Madagascar andMexico are examples referred to in the regional accounts.

Illegal tradeThere is no doubt that illegal trade in wild succulentplants continues in contravention of CITES (see Table

Akmrni//~k~ limonensis in cultivation at Can Te, A.C.gardens, Mexico.

2.5). Smuggling of small quantities of rare wild plants insuitcases or through the post, or mis-declaring of openlyimported shipments are examples of the activities whichcontinue and are extremely difficult to stop given thecurrent level of resources. Surveys conducted byTRAFFIC in Europe, Japan, and South Africa in the pastfew years have revealed the widespread and often openavailability of Appendix I listed wild plants in nurseries.The recent survey of European nurseries by TRAFFICEurope revealed that wild plants of the Appendix I cactiAriocarpus, Aztekium, Obregonia, Pelecyphora, andStrombocactus were all on sale in Italy. Austria has been asource of wild specimens of Ariocarpus on sale in Italy,but does not report in detail on its plant trade. WildAriocarpus plants have been sold openly in theAmsterdam flower market, and nurseries in Belgium andGermany continue to stock Appendix I rarities. Early in1992 seventy cacti including Ariocarpus and Strombocactuswere seized from a shop in Paris.

A clear-cut example of illegal international trade inrare cacti is provided by the commercial availability of therecently described species Aztekium hin tonii andGeohintonia mexicana. The original descriptions of thenew taxa were published in the Mexican journal Cactaceasy Suculentas Mexicanas at the end of 1991. Otherspecialist journals publicised the discovery of these newcacti (Geohintonia is a new genus). The following year thenew taxa were already present in commercial nurseriesand private collections within Europe. All specimensoutside Mexico are illegal both under Mexican nationallegislation and under CITES, as no export permits havebeen issued.

At the CITES Conference of the Parties, held inKyoto in 1992, Mexico called on importing countries tohelp with problems of illegal plant exports. In 1991, 18foreigners were caught illegally collecting cacti in Mexico;in some cases these were pseudo-conservationists “saving”

36

Geohintonia mexicana

wild plants. Recently, Mexican authorities have beenstudying where the collectors go to within the country andconcentrating enforcement efforts in those areas.

Trade in non-CITES succulents

International trade in non-CITES succulents involves awide range of species which frequently appear to bethreatened in the wild and/or collected in contraventionof national laws. This is the case, for example, withsouthern African succulents. A recent survey of Germanand Dutch succulent plant catalogues carried out for theGerman CITES Scientific Authority recorded theavailability of threatened southern African succulents at16 nurseries and noted indications of their wild status(Schippmann 1993). Although data on volumes of tradecannot be collected in this way, surveys of this kind can bea useful step in indicating species of potential concern.

In situ conservationSara Oldfield

Protecting wild plants in their natural habitats is generallyconsidered to offer the best long term chance of survivalfor rare and threatened species. In situ protection cantake various forms including legal protection of thehabitat where the species occurs; designation of micro-reserves, nature reserves, national parks or othercategories of protected area; land use controls or zoningrestrictions; protection on private land; voluntarymanagement agreements; habitat restoration; or thedevelopment of recovery plans for wild populations ofthreatened species. This section provides examples ofdifferent forms of in situ conservation for cacti and othersucculents to supplement the information within theregional accounts.

Protecting the habitats of threatened species

Areas specifically protected for threatened species arerelatively uncommon as most plant conservationlegislation is concerned with protecting species fromvarious forms of collection and disturbance. Habitatprotection for threatened plants is, however, included innational conservation legislation for several Europeancountries and also in the USA. The US EndangeredSpecies Act gives protection to the ‘critical habitats’ ofthreatened species including various cacti and othersucculents. Critical habitats are defined as areas which areessential to the conservation of the species concerned.These areas must be designated and their boundariesprecisely described in the Federal Register. As of October1987, of the 168 listed species of wild flora, there were 23species for which critical habitats had been designated(Groombridge 1992).

Protection of the habitats of threatened plant species

Habitat for Euphorbiacanariensis, LosPalmitos Valley, GranCanaria.

is also enshrined in regional conservation legislation. InEurope, for example, the EC Habitats Directive addressesthe conservation of both natural habitats and habitats ofthreatened species by the establishment of a Europeannetwork of Special Areas of Conservation (SACS) whichcollectively form the Natura 2000 network. New sites,identified by member states, in conjunction with the EC,include areas of natural habitat type listed in Annex I ofDirective, with certain habitats given special priority, andhabitats of the threatened species listed in Annex II.Habitat types included in Annex I which may beimportant for succulent plant protection includeMediterranean and pre-steppe brush (low formations ofEl~phorbia close to cliffs), various categories of rockyslopes, and Mediterranean montane forests.

Criteria for the selection of sites both at a nationallevel (Stage 1) and EC level (Stage 2) are given in AnnexIII of the Directive. The criteria for selection of sites forspecies include size, density, and isolation of thepopulation in relation to the national total, condition ofthe site, and global importance. All identified nationalsites of Annex II species will be considered as sites of ECImportance. Protection of the habitats of those endemicsucculents of the Canary Islands included in Annex II (seeTable 2.3) will therefore be protected under the Directive,reinforcing the protected area system outlined in Box 2.2.

The Habitats Directive obliges Member States toestablish conservation measures for SACS withappropriate management plans, and to avoid the

Box 2.2 Protected areas in the CanaryIslands

The Canary Islands have a number of protected areaswhich are important for succulent plant conservation.Examples of succulent species which are protected in situare included in Annex 6. Major National Parks occur onTenerife, La Palma, La Gomera, and Lanzarote. In contrast,Gran Canaria, the island which has experienced thegreatest degree of habitat destruction and modification,had, until recently, a relatively poorly developed series ofprotected areas. In the late 1980s a comprehensive reviewof species and habitats was undertaken on Gran Canaria’in order to select sites in need of protection. As a result ofthe review, PEPEN - A Special Plan for the Protection of theNatural Areas of Gran Canaria - was published in 1986.

The following year, the Regional Government passed anintermediate law putting most of the proposed areas ofPEPEN in a schedule of protected areas. It is now theresponsibility of the Direction General de Medio-Ambienteto prepare management plans for these sites. Despite theprogress in protecting the unique succulent plant flora ofthe Canary Islands, there are however, still significant gapsin the protected area coverage. Lowland plantcommunities are, for example, not adequately covered. Anew National Park is currently proposed to protect a largesector of the Euphorbia communities and eroded volcaniclandscapes of Gran Canaria.

Source: Synge 1991

deterioration and disturbance of the protected sites.Environmental Impact Assessments are required for anydevelopments which may adversely affect SACS.

Protected areas

The more common form of in situ protection for succulentplants is inclusion within the boundaries of land set asidefor general conservation purposes. A protected area isdefined as:

An area of land andlor sea especially dedicated to theprotection and maintenance of biological diversity, and of.natural and associated cultural resources, and managedthrough legal or other eflective means. (IUCN 1994a)

At a global level, a recent study of habitat protectioncarried out by WCMC indicates that overall 7.7 per centof tropical regions are conserved within protected areas.Protected area coverage of different habitat types isuneven with wet and moist major habitats betterrepresented in protected areas than dry major habitats.This probably reflects national and foreign policies topromote the conservation of tropical rain forests as wellas the fact that drier habitats are more prone toagricultural conversion (Green et al. 1995). The lack ofprotection for drier areas may indicate that succulentplant diversity is not adequately protected in the tropics.Green et al. (1995) note particularly the restrictedprotection for lowland dry and arid habitat types inCentral America, the Caribbean, and South America.The study also notes the need to carry out similarquantitative assessment of the representation of centresof species diversity, speciation, and endemism withinprotected areas.

For plants, the Centres of Plant Diversity (CPD)project coordinated by IUCN has identified some of themost important sites for species diversity and endemismworldwide. These are the global priority areas forconservation of plant biodiversity. Individual CPDs whichare particularly noteworthy for succulent plant species arelisted in Table 2.6, which also indicates the extent towhich the sites are currently protected. It is apparent thatmany CPDs are not legally protected, or are onlyprotected in part. A considerable proportion of those siteswhich are officially protected are not effectively managed(WWF and IUCN 1994).

International designations

At an international level various mechanisms exist toprotect globally important ecosystems and the speciesoccurring within them. One such mechanism is theConvention Concerning the Protection of the WorldCultural and Natural Heritage (World HeritageConvention). This Convention provides for thedesignation of areas of “outstanding universal heritage”as World Heritage sites, with the principal aim offostering international cooperation in safeguarding theseimportant sites. Few World Heritage sites are important

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Table 2.6 Centres of Plant Diversity - Conservation status of succulent rich sites

Site

Cal Madow, Somalia

Hobyo area, Somalia

Ogaden, Somalia /Ethiopia / Kenya

Socotra, Yemen

Cape Floristic Region,South Africa

Karoo-Namib Region,South Africa/Namibia

Madagascar

Canary Islands

Pinar del Rio, Cuba

Cockpit Country,Jamaica

Tehuacan Valley,Mexico

Edwards Plateau,Texas, USA

California FloristicProvince

Atacama Desert, Chile

Lomas, Peru/Chile

Vegetation

Dry montane forest

Deciduous bushland, woodland,dunes

Deciduous bushland, woodland

Semi-desert, shrubland, thicket,grassland

Fynbos, shrubland, montane

Succulent shrubland

Predominantly forests and thicket

Coastal vegetation, Euphorbia-richvegetation, scrub, woodland, forest

Seasonal forests, succulent andthorn scrub

Subtropical forest, scrub thicket

Dry scrub, deciduous forest

Forest, grassland, semi-desertscrub

Forests, woodlands, coastal scrub,grasslands

Mixture of annual, short-livedperennial and woody scrub

Islands of low montane desertscrub and thorny steppe

for succulent plant conservation. Exceptions are theReserve naturelle integrale du Tsingy de Bemaraha inMadagascar inscribed as a World Heritage site in 1990,the Grand Canyon National Park in Arizona, USA, andthe Galapagos Islands.

Biosphere reserves are created under the UNESCOMan and the Biosphere (MAB) Programme. Theobjective of this Programme is to develop a scientific basislinking the natural and social sciences for the rational useof and conservation of the biosphere and for theimprovement of the relationship between humans andtheir environment. Biosphere reserves are multipurposeareas dedicated both to the conservation of characteristicecosystems and species, and to sustainable developmentto meet human needs. Currently the internationalnetwork consists of over 285 biosphere reserves.

Conservation status

Daalo Forest Reserve, no protection in practice.

No protected areas, extremely vulnerable to overgrazing.

No protected areas.

No protection, but traditional practices have preventedserious exploitation.

Reserves cover 19% of region but mainly in mountains.

National Park and a few other reserves covering about 2%of region.

Protected areas cover less than 2% of the island.

Many protected areas with some notable gaps forsucculent rich sites.

National Park; lowlands at risk.

No protected areas.

Small botanic garden.

Protected areas cover less than 0.5% .

About 11% of land protected, mostly montane, lowlandhabitats threatened.

Two protected areas cover 468 km*, many endemics needmore extensive protection.

Lachay Nature Reserve, protected area coverageinadequate.

Various biosphere reserves are particularly important.for succulent plant conservation. Mexico has sixinternationally recognised biosphere reserves includingthe Pinacate Biosphere Reserve covering 480,000 ha ofthe Sonoran Desert and adjoining the Organ PipeNational Monument in the USA (see below). The MapimiBiosphere Reserve, established in 1977, covers over100,000 ha of Chihuahuan Desert with more than 20species of cacti (Box 2.3).

In the USA, an area of succulent plant diversity isprotected in the Big Bend National Park, which is one of47 biosphere reserves in the country. This National Parkprotects a large area of Chihuahuan Desert in BrewsterCounty, Texas. Rare cacti occurring within Big Bendinclude Ancistrocactus tobuschii, Coqphantha ramillosa,Epithelantha micromeris var. bokei, and Sclerocactusmariposensis.

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Box 2.3 Reserva de la Biosfera de Mapimi, M6xico

Location Situated north-east of the town of Ceballos on the boundaries of the states of Durango, Chihuahua, and Coahuila.The reserve lies in a hollow or basin surrounded by small mountain ranges running more or less parallel from north to south, inthe large catchment area known as Bolson de Mapimi.

Area 103,000 ha (expanded from original 100,000 ha; core area 38,000 ha)

Land tenure There are various common public lands, private ranches, and small peasant properties.

Physical features The reserve is part of the riverine basin (endorheic) system of the Mapimi Bolson of the North MexicanCentral Tableland and part of the Chihuahuan Desert. Alluvial deposits predominate, consisting of recent Pleistocene gravels,clays and muds, Outcrops of igneous rock from the Tertiary period and volcanic rocks (rhyolites, andesites, and basalts) arealso found. The landscape consists of isolated mountains and extensive interconnected plains.

Vegetation The reserve is located in the Chihuahua arid zone, typically represented by a restricted flora with a large number ofendemics. There are five vegetation types, all generally open and xerophytic. Microphyllous matorral scrub is the mostrepresentative vegetation community, as characterised by Larrea divas-&fa, Agave spp., Hechtia spp., Pasfinaca spp., Euphorbiaantisyphilitica, and Opuntia spp.

Local human population In the early 1980s there was a total population of approximately 100 people. Of the main ranch andfarm units in the reserve at this time, three were privately owned while eight were ejidos in which the land was entrusted by thegovernment to a local community. Nine of the units were devoted to stockraising, one to the extraction of wax from candelilla,whilst 11 extracted salt from a lagoon to the north of the reserve. Agricultural development is limited.

Scientific research and facilities Research facilities include the Desert Laboratory, established in 1978 by the lnstituto deEcologia and located on 20 ha of land in the centre of the reserve. Research includes studies on amphibians and reptiles, raptorbiology, the regeneration of desert vegetation after over-grazing, fire, and various types of human activity.

Conservation management Established primarily to protect the Bolson tortoise and the fragile arid habitat. When the reservewas first established there was no zonation, but a core and buffer zone system has subsequently been established. Thescientific management of the biosphere reserve is coordinated by the lnstituto de Ecologia. The local people are involved in alegally constituted association to assist in management of the reserve. This group includes representatives of the cattle ranches,small land owners ejidos, the National Council of Science and Technology (CONACYT), and the lnstituto de Ecologia.

Source: WCMC Protected Areas Database

National designations role in the conservation of biodiversity worldwide-Protected areas, defined as sites with an area of 1,000 ha

(Groombridge 1992). In general, protected areas

or more in IUCN categories I-V (see Box 2.4), have beendesignated by national legislation have been developed to

established in over 160 countries and play an essentialprotect scenic landscapes, representative ecosystems, or

Box 2.4 IUCN Categories of protected areas (I-V) with their management objectives

Scientific Reserve/Strict Nature Reserve: to protect nature and maintain natural processes in an undisturbed state inorder to have ecologically representative examples of the natural environment available for scientific study,environmental monitoring, education and for the maintenance of genetic resources in a dynamic and evolutionary state.

II National Park: to protect natural and scenic areas of national or international significance for scientific, educational andrecreational use.

III /Vatura/ Monument/Natural Landmark: to protect and preserve nationally significant natural features because of theirspecial interest or unique characteristics.

IV Managed Nature Reserve/Wildlife Sanctuary: to assure the natural conditions necessary to protect nationally significantspecies, groups of species, biotic communities, or physical features of the environment where these require specifichuman manipulation for their perpetuation.

V Protected Landscape: to maintain nationally significant natural landscapes which are characteristic of the harmoniousinteraction of man and land while providing opportunities for public enjoyment through recreation and tourism withinthe normal life style and economic activity of these areas.

Source: KJCIV (1994a)

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animal populations rather than to protect individual plantspecies or botanical diversity. There are a few notableexceptions. Two protected areas in Arizona, USA, havebeen designated specifically to protect cacti. These are theOrgan Pipe Cactus and Saguaro National Monuments.The former of the two protects an important example ofSonoran desert vegetation with, in addition to the organpipe cactus Lemaireocereus thurberi, 28 other taxa of cactioccurring within the protected area.

In many other countries succulent plants occur withinprotected areas designated for general conservationpurposes (see Box 2.2) but often full botanicalinventories have not been compiled for these sites. Oftenit is unclear which succulent species occur within aparticular protected area, and also to what extentparticular rare or threatened species are actuallyprotected in situ in various parts of their range.

At a national level, some countries have not yetdeveloped protected area networks, and in othersimportant areas of succulent-rich vegetation are not yetadequately represented. Most of the countries reviewed inthe Action Plan have important areas of succulent plantdiversity which are not yet included in protected areasystems. These include Brazil, Madagascar, Mexico,Caribbean islands, Somalia, South Africa, and Zimbabwe.Recommendations for priority sites for protected areasare given in Chapter 4.

With increasing threats to succulent species in the wildand increasing land demands, implementation ofrecommendations for in situ protection will be particularlychallenging. Novel approaches may be required. One suchapproach is the creation of micro-reserves specificallydesigned to protect endangered or endemic plant species.An example of this approach is being pioneered forendemic species in Valencia, Spain. In 1994, the RegionalGovernment of Valencia created a new kind of legalprotection, with the aim of establishing small permanentreserves for plants in need of habitat protection (Laguna1997). More than 60 small reserves have already beencreated, with an individual area of less than 20 ha, and inthe next few years a total of 150 micro-reserves will beestablished protecting 280 endemic species. The small sizeof each protected area is expected to reduce the risks ofproblems with land users. Management of the micro-reserves in Valencia is coordinated with the developmentof recovery plans and ex situ conservation measures. It isclearly resource and management intensive and will notbe appropriate in all succulent rich areas requiringprotection. Such an approach may however beappropriate, for example, for narrow endemics in parts ofthe Caribbean, Madagascar, Mexico, and South Africa.

The increasing availability of distribution data forsucculent plant species and computerised mappingsystems make it easier, in theory, to select and designareas for the protection of succulent diversity, althoughthe actual designation of national parks and reservesremains challenging because of conflicting landuse

demands. The use of geographic data from literatiu-e,herbarium specimens, and other sources to p lan for theconservation of plant genetic resources is discussed byMaxted et al. (1995). Similar techniques can be used toplan conservation strategies and protected areas for rareand threatened plant species.

In Mexico, the distributions of endangered cacti in theChihuahuan Desert Region have been mapped on a gridsquare basis to show the areas of maximum diversity ofthe species (Hernhndez and Barcenas 1995). In this waythe areas of particular importance for cactus conservationare identified, as discussed further in the MexicanRegional Account in Chapter 3. Other factors would needto be taken into account in the selection of formallyprotected areas such as habitat diversity, presence anddiversity of other species groups, and land use pressures.However, the study of the Chihuahuan Desert Regionvividly highlights the fact that none of the criticallyimportant species rich quadrants identified forendangered cacti are included in protected areas.

Increasingly it is recognised that designation andmanagement of protected areas must take into accountthe views and requirements of local people. TheRichtersveld National Park in South Africa, brieflydescribed in Box 2.5, was established in 199 1 followingnearly 20 years of consultation with local people.

Box 2.5 Richtersveld National Park,South Africa

The Richtersveld National Park in the north-western Capeof South Africa protects an internationally important area ofsucculent plant diversity. The protected area wasdesignated in 1991 after extensive local consultation.Management involves recognition of local needs with localcommunities involved in the decision-making process. ThePark covers 1,624 km2 of attractive desert scenery andunique endemic vegetation. It is particularly important forthe conservation of South Africa’s mesembs, together withdistinctive Aloe pillansii, A. dichotoma, A. qeyeri, andPachypodium namaquanum. Threats to the’succulentplants have included mining, stock farming, and collectionfor horticulture. Tourism is now a growth industry and thisin itself places pressures on the succulent plant species.

Private land

Acquisition and management of land for conservation byNGOs and individual land owners are important means ofensuring in situ conservation. In the USA, for example,The Nature Conservancy Council administers over 1300reserves covering 650,000 ha; the National AudubonSociety owns or leases over 100 sanctuaries covering over60,000 ha; and Operation Stronghold is an alliance of 800-900 private landowners who have undertakenconservation measures on private land estimated to cover2-2.5 million ha (WCMC 1992).

In Mexico, the NGO Can Te, A.C. has a programme,‘Comprar para Conservar’ or ‘Purchase to Preserve’,

41

which encourages people to contribute financially towardthe purchase of threatened habitats. Can Te, A.C.considers that most threatened species of cacti and othersucculents do not grow in areas appropriate for nationalprotection. In these cases, purchase of the land may bethe only viable option to preserve the sites. Realistically,this can be best achieved by non-governmentalorganizations or by private individuals. Can Te, A.C. haspurchased the terrain of Mammillaria albiflora on abarren hillside in north-eastern Guanajuato. Collectorshave not appreciably reduced the population, even thoughit is the main known locality for the taxon. Recently, theplant was put in jeopardy when the owner of the propertyoffered it for sale for agricultural use. However,purchasing land is not always practical. For example, theexpansion of the city of Queretaro threatens to eliminatethe only known remaining habitat of Mammillariamathizdae, a cactus little in demand and not threatened bycollectors. An attempt by Can Te, A.C., in cooperationwith the Queretaro campus of the Instituto Tecnologico yEstudios Superiores de Monterrey (ITESM), to purchasethe habitat was unsuccessful due to the high pricesdemanded by the speculating owners.

Species management

Within areas set aside for conservation, management ofthe plant communities will frequently be required.Protected areas generally have management plans whichset out the requirements for maintenance of the site inbroad terms. For populations of individual rare orthreatened species, particular management prescriptionsmay be necessary. As Given (1994) points out, “Particularspecies may be threatened or in decline because of factorsthat are independent of the ecosystem as a whole.Examples include harvesting, predation and the

Box 2.6 Recovery plan for Pediocactusknowltonii

Pediocactus knowlfonii is an Endangered cactus of theUSA, with only one viable population on a single hill, southof La Boca in northern New Mexico. The species was listedas Endangered under the Endangered Species Act in 1979when only about 1000 individuals were known to remain.The decline of the naturally rare species has been causedby habitat loss, collection for horticulture and, in 1960, amisguided “rescue” operation when thousands of plantswere collected from the proposed site of a dam. TheKnow/ton Cactus Recovery Plan approved on 29 March1995, called for restoration of the La Boca population toapproximately 100,000 cacti, long-term protection of thesite, and reintroduction of the species at other suitablelocalities, The site is owned by The Nature Conservancy. Ithas been fenced to keep out cattle and in an attempt todeter collectors. The development of detailed recoverygoals have been hindered by lack of biological data for thespecies.

Source: Endangered Species Technical Bulletin IO (12).1995.

consequences of small population size. Some species mayrequire specific habitat manipulation independent of thatapplied to the whole system where they occur.”

Species management within protected areas mayinvolve protection from unwanted human disturbance,with periodic monitoring or more active manipulation.Unfortunately knowledge of the biology and ecology ofsucculent plants species is frequently very limited, andmanagement where it is applied is likely to be based ontrial and error. Research from ex situ situations can beparticularly helpful for succulent plants. Conservation ofsucculent plant species in the long-term will be mostsuccessfully achieved by an integrated approach involvingboth in situ and ex situ conservation measures.

Ex situ conservationEdward Anderson

As seen in the introductory chapters of this Action Plan,many succulent taxa are in serious trouble, and may belost unless immediate conservation efforts areundertaken. Cultivation of plants in botanic gardens,arboreta, private collections, and nurseries is an ex situconservation method that is used by biologists to assure,at least, some protection of species ‘away from danger’.Plant material in ex situ cultivation may be used forreinforcing existing wild populations or in the re-establishment of wild populations following the IUCNGuidelines for Re-Introductions (IUCN 1995).

In 1985 an international conference was held in LasPalmas de Gran Canaria, in which the organisationspresent adopted a series of 13 recommendations dealingwith ‘Botanic Gardens and the World ConservationStrategy.’ Recommendation 4 dealt specifically with exsitu conservation. It stated (IUCN 1986):

“While recognising that no single approach to theconservation of endangered species can be relied upon;appreciating that ex situ conservation is a necessaryadjunct to in situ conservation; and acknowledging theimportance of seed banks in the long-term conservationof genetic resources, The International Conference onBotanic Gardens and the World Conservation Strategy:

Urges Botanic G a r d e n s t o recognise the i rresponsibility to maintain, propagate and makeavailable stock of critically threatened species forscientific and horticultural research, for reintroduction(where appropriate) and to provide suitable stock forhorticulture;Recommends that exploration and collection ofspecies be based on concepts of infraspecific diversityso that ecogeographical diversity and diversitybetween and within populations be sampled in such away that a maximum of genetic diversity be capturedand stored;

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Urges Botanic Gardens to become involved with seedconservation and recommends that the InternationalBoard for Plant Genetic Resources (IBPGR) (now theInternational Plant Genetic Resources Institute -IPGRI) be approached so as to establish closercollaboration on the conservation of threatenedspecies;Recommends that every effort be made to maintainminimal international standards for seed storage andrejuvenation;Recommends IUCN to continue and expand themonitoring and co-ordination of ex situ conservation,presently carried out by the Botanic GardensConservation Co-ordinating Body, as an integral partof the implementation of the Botanic GardensConservation Strategy;Recommends Botanic Gardens and other relevantinstitutions to support this essential work and toprovide the necessary finance for it.”

As a response in part to these recommendations andbecause of the important succulent collections that existaround the world that could contribute significantly to theconservation of succulents, this section deals with variousforms of ex situ conservation presently available forsucculents. It also describes several actions that might betaken to facilitate ex situ conservation.

Conservation collections have been defined as “livingcollections of rare or endangered organisms, establishedfor the purpose of contributing to the survival andrecovery of a species” (Center for Plant Conservation1991). There are four important types of succulent plantcollections that may be utilised in ex situ conservation

botanic gardens and arboreta, seed banks,activities:private collections, and, to some extent, commercialnurseries. Each of these plays a significant, thoughdifferent role in conservation. Nonetheless, carefulnetworking among these different collections and theirmanagers or owners will provide botanists andconservationists with an effective tool for the long-term

Aeonium tabuliforme, a Rare Crassulaceae of theCanary Islands.

preservation of germplasm of rare and endangeredsucculents. Additionally, plant material in c’x sitzrcultivation may be used for reinforcing existing wildpopulations or in the re-establishment of wild populationsfollowing the IUCN Guidelines for Re-in troductions(IUCN 1995).

Botanic garden networks

The International Association of Botanic Gardens(IABG) is an umbrella organisation for all the 1600botanic gardens and arboreta worldwide. Some arc thetraditional botanic gardens, whereas others are naturereserves, small private gardens, nurseries, and agriculturalexperimental stations. These institutions combinedprovide more than 100,000 hectares (247,100 acres) of PXsitu habitat for the cultivation of plants. There are alsoregional associations of IABG in Latin America and theCaribbean, Asia, and Europe. In North America theAmerican Association of Botanical Gardens and Arboretais also active.

Following the recommendations of the Las Palmasconference, IUCN established the Botanic GardensConservation Secretariat, BGCS, (which later becameBotanic Gardens Conservation International - BGCI), toco-ordinate botanic gardens’ conservation workworldwide. In 1985 the Secretariat published two reportsof significance to succulent plant conservation: ReportNo.13 of the Rare and Threatened Species of MexicanCacti and Report No.15 of the Succulents of Africa andMadagascar. The Botanic Gardens Conservation Strategywas published in 1989 (IUCN-BGCS 1989) and provides arationale and basic guidelines for plant conservation workthrough botanic gardens. Today over 450 gardens andother institutions from 90 countries are members, andBGCI has an active programme to enhance theconservation and environmental education efforts ofbotanic gardens worldwide. BGCI maintains a database ofrare and endangered species currently (1995) holdingabout 200,000 records of plants in botanic gardens, manyof which are succulents.

In 1993 the IABG and BGCI signed a wide-rangingagreement “for the purposes of fostering the developmentof the world network of botanic gardens and arboreta,towards researching, documenting, conserving andutilising the plant resources of the world. . .” (BotanicGardens Conservation News 1994).

The World Conservation Monitoring Centre(WCMC), in Cambridge, UK, maintains a database ofover 100,000 rare and threatened plant taxa worldwide.Conservation biologists, particularly members of theIUCN/SSC Specialist Groups, provide data to WCMC,which in turn compiles, analyses, and supplies valuc-added information regularly to Botanic GardensConservation International (BGCI), the IUCN SpeciesSurvival Commission, and many other international andnational organisations and institutions involved inmonitoring rare plants.

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In the United States the Center for PlantConservation (CPC) was founded in 1984 at the ArnoldArboretum. Currently headquartered at the MissouriBotanical Garden, the Center is a consortium of botanicgardens and arboreta in the USA whose purpose is topreserve critically endangered plant taxa of the USA byestablishing a national ex situ collection of rare plantgermplasm. This National Collection is held in theCenter’s 25 participating institutions across the country,and is primarily composed of stored seeds which havebeen collected in a manner to capture the geneticdiversity of a taxon to the fullest extent possible. Thecollection, and others like it, serves several functions: 1)as an insurance policy against extinction in case of acatastrophic loss in the wild; 2) as a source of material forresearch and education programs which can contribute tothe conservation of a species; and 3) as a source ofgermplasm for potential re-introduction and ecologicalrestoration projects, which are carried out on acooperative basis.

Over 450 plants are currently represented in theNational Collection. Of these, approximately 25 speciesare succulents, with the majority at the Desert BotanicalGarden in Phoenix, Arizona. Genetically representativeseed collections of the rare succulents have been placed inlong term storage at the Desert Botanical Garden,Ranch0 Santa Ana Botanic Garden (California), SanAntonio Botanical Garden (Texas), and FairchildTropical Garden (Florida). Research is conducted ongermination and viability testing, propagation methods,and reproductive biology. These botanic gardens are ofteninvolved in surveying, monitoring, and re-introductionprojects with federal and state agencies, as well as otherpartners.

National networks of botanic gardens exist in manyother countries. For example, the Australian Network forPlant Conservation (ANPC) has established a programmein ex situ conservation demonstrating how botanic gardenscan network effectively in conservation matters. AnEndangered Species Collection was started in 1986 at theAustralian National Botanic Gardens, Canberra, forresearch, education, display, and possible re-introduction.This collection possibly will become available to thenursery industry as well (Richardson 1992). In theNetherlands, the Dutch Botanic Garden Foundation co-ordinates a Decentralised National Collection amongstthe Dutch botanic gardens, including very significantnumbers of cacti and other succulents. Active botanicgarden networks involved in conservation also operate inmany countries, including Brazil, China, Colombia, Cuba,France, Germany, Indonesia, Italy, Japan, Mexico,Russia, Spain, and Portugal (Wyse Jackson 1993).

DatabasesThe development of the International Transfer Format forBotanic Garden Plant Records (IUCN-BGCS 1987) andcomputer-based plant collection software systems have

provided the means for, and been influential in, thedevelopment and improvement of plant records andcollection documentation worldwide.

One of the mos t se r ious p rob lems fac ingconservationists is that of nomenclature. Some raresucculents are referred to by a variety of names; forexample, the Office of Scientific Authority of the US Fishand Wildlife Service and the CPC continue to use thenomenclature of Benson (1982) for cacti in the USA,although CITES has now published an updated checklistof the most commonly accepted names of cacti based onrecent research (Hunt 1992). In an effort to assistscientists and horticulturists in dealing with the perplexingproblems on nomenclature, the contents of all issues ofthe Repertorium Plantarum Succulentarurn (RPS) havebeen compiled by the Stadtische Sukkulenten-SammlungZurich (ZSS), creating the RPSIZSS Database. Not onlydoes this database contain thousands of names, but it alsocontains curatorial information as needed for living andherbarium collections. The following custom-designedprintouts are available: taxonomic data by family,taxonomic data by genus, and synonymy of a genus. Suchsynonymy lists are available for most succulent families.Additional lists are published in the annual issues of theIOS Bulletin.

CollectingThe CPC (1991) has published guidelines for collectors ofrare plants in response to some concern about inadequategenetic representation of endangered species in cx situcollections. The guidelines address five questions whichconstitute a natural hierarchy covering species,populations (and ecotypes), individuals, and alleles. Thework of the collector may also be influenced by the degreeof genetic difference among populations, time and moneyconsiderations, the survival rate of propagules, and theintended use of the collections. The Center’s fivesampling questions are:

Which species should be collected? The two mostimportant criteria here are the probability of loss of aunique genepool, and the potential for restoration orrecovery.How many populations should be sampled per species?As a rule, one to five ecotypes or populations shouldbe sampled, depending on the population history andthe degree of difference among the groups.How many individuals should be sampled perpopuZa tion ? Ten to fifty individuals should be sampledper population, but this depends on population sizeand genetic mobility.How many propagules should be collected from eachindividual? From one to twenty should be obtained,depending on the survival rate of the propagules andthe long-term needs of the collection.Under what circumstances is a multiyear collection planindicated? More than one year is recommended,

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especially if the desired number of propagules cannotbe removed safely without affecting the reproductivecapabilities or demography.

Developing techniques, particularly for DNA study,for the analysis of genetic variability of plant populationsin the wild and maintained ex situ provide valuable meansfor improving the genetic representation in gardencollections. Such techniques are also becoming useful forconservation, to study material from garden sources (frombotanic gardens, private growers, and commercialsources) that has lost its documentation.

Seed banks

There is an increasing number of seed banks beingdeveloped for wild plants worldwide, particularly bybotanic gardens, although there is still relatively little co-ordination between them. A 1994 survey by BGCIindicated that there are currently over 150 such seedbanks in botanic gardens. The extent to which succulentspecies are contained within their collections is unknown,however. One such collection specialising in the storage ofarid land species is held at Wakehurst in the UK, and asecond by members of the CPC in the US where seedsfrom the National Collection of endangered plants aredesiccated and frozen for long term storage of thegermplasm.

The collection and ex situ storage of seeds of rareplants present several significant conservation problems.The first concerns the impact of collecting seeds on thenatural populations. This is of special concern withrespect to succulents occurring in arid regions, for thesevere climatic fluctuations in deserts frequently lead tosporadic reproduction. Thus, collection of seeds fromplants with low levels of reproduction during periods ofstress can be highly detrimental to the long term survivalof that population. IUCN-BGCS (1989) recommends thatnever more that 10 per cent of the seed available in a wildplant population should be collected.

A second concern, expressed by Richardson (1994)involves the exchange of seeds in Index Seminumprograms in which weedy plants are distributed withinsufficient controls. Thus, exchange programs by botanicgardens with good intentions may result in theintroduction of potentially dangerous exotic plants. TheCPC has recognised the problems, both potential andreal, of seed banks and the exchange of seeds, and haspublished extensive guidelines for the management of“orthodox” seeds (those that tolerate severe desiccationor dehydration and which can usually be stored for longperiods of time) (Wieland 1995). These guidelines dealwith the collection, storage, and shipment of seeds. Muchinformation on the management of seeds has also beenpublished by IPGRI.

Seed collections areconse rvation, for they pother than from the field

of ex situermplasmguidelines

are followed, seed banks and Index Seminum exchangescan play an important role in perpetuating raresucculents, as well as providing research and horticulturalmaterials. The National Botanical Institute of SouthAfrica (NBI), including its network of botanic gardens hasa seed exchange programme with other institutions; seedsare exchanged “on the understanding that it is not usedcommercially.” These seed collections can also bevaluable to some in situ conservation projects whichinvolve re-introduction of plants into habitat. Thoseorganisms placed in habitat may have originated fromseeds propagated in either botanic gardens or commercialnurseries. Guidelines for such plant re-introductions havebeen produced by several organisations, including theIUCN-SSC Re-introductions Specialist Group (IUCN1987 and 1995), and the Handbookfor Botanic Gardens onthe Re-introduction of Plants to the Wild has beenpublished by BGCI (Akeroyd and Wyse Jackson 1995).

Botanic gardens

A wide range of botanic gardens maintain very extensivecollections of succulent plants. Compared to many otherplant groups, the rare and endangered species of cacti andsucculents are well represented in many collections. A fewof these gardens grow collections of the succulentmembers of their own flora, e.g. the Jardin Bot5nicoCanario ‘Viera y Clavijo’, Gran Canaria, Spain; theNational Botanic Institute, South Africa; and the KingsPark and Botanic Garden, Perth, Western Australia.However, the succulent collections of most botanicgardens are of exotics. There are also many regions ofnatural succulent vegetation, such as Ethiopia, Somalia,Chile, where there are no functioning ex situ conservationfacilities. The development of local facilities and expertiseshould be emphasised in plans to establish newcollections. Collaborative training programmes betweenthe larger international botanic gardens and local facilitiesshould occur.

The following eight examples are from botanicgardens that have already shown leadership in managingcollections of succulents, and are actively involved inconservation work.

1. University of California Botanical Garden U CBerkeley, Centennial Drive, Berkeley, California 94720,USA. Telephone: (510) 6438040. FAX: (510) 642-5045.

Founded in 1890, this state supported garden is part ofthe largest university campus in the USA. The garden,located in Sycamore Canyon above the main campus, hasan area of 13 hectares (32 acres). The succulent collectioncomprises a fifth of the total collection. There are 2570taxa, comprising 4046 accessions, with 2293 species of thefamilies Agavaceae, Aizoaceae, Cactaceae, Crassulaceae,and Aloe (included in Liliaceae in their data base). Ofthese, 69 per cent of the taxa are documented (78 per centat the species level). The Berkeley garden also has livingmaterial of 45 species of CITES Appendix I succulents. It

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is a participating member of the CPC. Living material ofthe following collectors of succulents are present in thegarden: M. Cardenas, S. B. Hogan, P. C. Hutchison, A.Lau, R. Moran, M. Kimnach, W. Krahn, F. Ritter, R.Rodin, and J. West.

2. Desert Botanical Garden 1201 North Galvin Parkway,Phoenix, Arizona 85008-3490, USA. Telephone: (602)941-1225. FAX: (602) 481-8124.

This garden is a private institution and was founded in1937. It covers an area of 53 hectares (145 acres), withmost specimens planted outside. There are also threeglasshouses and an extensive shade house. The Garden’ssucculent collection contains the following specimens:

Family No. taxa No. taxadocumented

AgavaceaeAizoaceaeAloaceaeAsclepiadaceaeCactaceaeCrassulaceaeDidiereaceae,EuphorbiaceaeFouquieriaceae

267 204 (76%)274 205 (75%)253 117 (46%)88 65 (74%)1296 841 (65%)102 55 (54%)13 1 (8%)124 68 (55%)14 8 (57%)

There are 38 species of CITES Appendix I succulentsin the collection in the Agavaceae, Apocynaceae,Cactaceae, Euphorbiaceae, Fouquieriaceae, andAloaceae. Significant collectors at the Desert BotanicalGarden have included E. F. Anderson, M. Baker, R.Engard, H. S. Gentry, W. Hodgson, P. C. Hutchison, F.Kattermann, G. Lindsay, B. Parfitt, D. Pinkava, J.Rebman, L. Slauson (Ecker), and A. Zimmerman.

The Desert Botanical Garden is a member of theCPC, and is responsible for protecting many of thethreatened and endangered succulents of the south-western USA. The Garden has established a seed bankand living ex situ collections of the following succulents:

Agave arizonicaA. parvifloraCoryphantha recurvataC. ramillosaC. scheeri var. robustispinaEchinocactus horizonthalonius var. nicholiiEchinocereus chisoensisE. viridiflorus var. davisiiEpithelantha bokeiEscobaria minimaE. robbinsorumE. sneedii var. sneediiMammillaria thornberiPeniocereus greggi var. transmontanusSclerocactus erectrocentrus var. acunensis

Mammillaria and Coryphantha collection, DesertBotanical Garden, Arizona.

S. erectrocentrus var. erectrocentrusS. mariposensis

The Garden has also established permanent in situmonitoring sites and is carrying out extensive field studiesfor several of the above taxa.

3. Huntington Botanical Gardens The Huntington, 115 1Oxford Road, San Marino, California 91108, USA.Telephone: (818) 4052160. FAX: (818) 4050225

The Huntington Botanical Gardens, founded in 1919,has a succulent plant collection, as well as other majorplant collections and an important arts and humanitiescollection and library. Plant records were begun in 1930.The desert plant collection covers an area of 6 hectares(15 acres). Most of the collection is outdoors, but thegarden also has several glasshouses. The succulentcollection consists of nearly 20,000 accessions,representing more than 8000 taxa, of which one-third aredocumented. Some of the most important succulents inthe collection consist of:

Family No. taxa No. taxadocumented

Agavaceae 490 304 (62%)Aizoaceae 728 255 (35%)Aloaceae 780 304 (39%)Asclepiadaceae 429 249 (58%)Asteraceae 74 34 (46%)Bromeliaceae 196 53 (27%)Cactaceae 3125 812 (26%)Crassulaceae 1293 427 (33%)Didiereaceae 11 1 (1%)Euphorbiaceae 483 97 (20%)Fouquieriaceae 13 8 (62%)

The Huntington has 60 CITES Appendix I taxa in itsliving collection. It is responsible for growing anddistributing the International Succulent Institute (ISI)

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materials each year. The list of succulents offered ispublished annually in the Cactus and Succulent JournalFJv*

Succulent collectors include E. F. Anderson, W.Baker, G. Barad, J. Bauml, J. Berdach, J. Betzler, J.Bleck, F. Boutin, S. Brack, F. Brandt, M. Cardenas, J.Clements, S. Collenette, J. Dice, J. Dodson, H. Earle, U.Eggli, A. Ellert, C. Fleming, J. Folsom, R. Foster, E. Gay,H. S. Gentry, C. Glass, S. Hammer, D. Hardy, A.Hoffman, I. Hoffmann, F. Horwood, P. Hutchison, E. vanJaarsfeld, F. Katterman, R. Kiesling, M. Kimnach, K.Knize, D. Koutnik, A. Lau, J. Lavranos, B. Leuenberger,S. Linden, H. Y. Liu, J. Lomeli-Sencion, G. Lyons, T.Mcdougall, N. Martinez, W. Minnich, R. Moran, L.Newton, F. Otero, D. Plowes, W. Rauh, H. Sanchez-Mejorada, R. Thompson, J. Trager, V. Turecek, C. Uhl,M. Vassar, and M. Wilkins.

4. Jardin Bothnico de1 Instituto de Biologia de UNAMInstituto de Biologia, Universidad National Autonoma deMexico. Ciudad Universitaria, Apartado Postal 70-614,04510 Coyoacan, Mexico D.F., Mexico. Telephone: 52-915-6-22-9046, 6-16-1297. FAX: 52-915- 6-22-9046, 6-16-2326.

This major Mexican botanic garden is part of theNational University of Mexico and is funded by thegovernment. The botanic garden consists of an area of 10hectares (24.7 acres). A major portion of the garden isdedicated to Dra. Helia Bravo H. and contains thesucculent collection. It consists of 148 Agavaceae species,with 120 (81%) documented (1 species of CITESAppendix I); 455 Cactaceae species, with 300 (66%)documented (23 species of CITES Appendix I); 65Crassulaceae species, with 60 (92%) documented; 25Nolinaceae species, with 15 (60%) documented.Important collectors of succulents have been H. Bravo H.,A. Garcia M. (active), D. B. Gold, U. Guzman C. (active),E. Matuda, F. Miranda, J. Reyes S. (active), H. Sanchez-Mejorada R., and S. Arias M. (active).

5. Jardin Botanic0 “El Charco de1 Ingenio” Can Te, A.C,Mesones 71, 37700, San Miguel de Allende, Guanajuato,Mexico. Telephone: 52-415-2-2990. FAX: 52-515-2-4015.

This private garden was founded in 1990 in an areanorth-west of the colonial city of San Miguel de Allende.At present much of El Charco is in the form of a naturereserve, but an area of 64 hectares (158 acres) is beingdeveloped into a more formal botanic garden. This newgarden is emphasising the Agavaceae, Cactaceae, andCrassulaceae of the Mexican flora. Most of the collectionis outside, but there are two large glasshouses on thegrounds of the garden and a major propagation facility ona farm a few kilometres away. Significant collectors are C.Glass and W. A. Fitz Maurice. A seed exchangeprogramme with other institutions has not yet beenformally approved by the Mexican government, but it isthe intention of Can Te, A.C. to develop a major seedexchange programme for the Mexican government.

Hopefully, this will reduce the pressure on wildpopulations from illegal collectors.

Can Te, A.C. has also been involved in several rescueand salvage operations in Mexico. Mammillai-iaaurilanata, Pelecyphora aselliformis, and Echirzocactusgrusonii are examples of cacti which have been removedprior to major habitat changes, such as road and damconstruction. Can Te, A.C. has also established a naturereserve for the protection of Mammillaria a1bif7ora. Thestaff is also heavily involved in rare cactus monitoringprojects in many parts of Mexico.

6. Jardin Exotique B.P.105, Monte Carlo 98002, Monaco.Telephone: 33-93-30-33-65. FAX: 33-93-30-60-74.

Founded 1933, this municipally operated garden, withan area of one hectare (2.5 acres), has long specialised insucculents. It contains 4000 succulent plant taxa, of which800 (20%) are documented. The collection contains about30 CITES Appendix I taxa in Pachypodium, Euphor-hia,Aloe, Ariocarpus, Astrophytum, Discocactus, Mammillaria,Melocactus, Obregonia, Pediocactus, Turhinicarpus, andUebelmannia. The most important succulent groupsrepresented are the Agavaceae, Aizoaceae, Aloaceae,Amaryllidaceae, Apocynaceac, Asclepiadaceae,Asteraceae, Boraginaceae, Bromeliaceae, Cactaceae,Crassulaceae, Dracaenaceae, and Euphorbiaceae.Collectors of succulents who have deposited material inthe Jardin Exotique include M. Kroenlein, W. Rauh, R.Kiesling, and C. Backeberg.

7. National Botanic Gardens of South Africa NationalBotanic Institute, Kirstenbosch, Private Bag X7,Claremont 7735, South Africa. Telephone: 27-2 1-797-2090. FAX: 27-021-797-2376.

The National Botanical Institute of South Africa haseight National Botanical Gardens strategically located inthe major natural regions of the country: Harold Porter,Karoo, Kirstenbosch, Lowveld, Natal, Orange Free State,Pretoria, and Witwatersrand National Botanical Gardens.The role of these gardens is to cultivate, protect, display,research, and utilise South Africa’s floral wealth, for theeducation and enjoyment of all. While all of these gardenshave succulent plants in their collections, three of thegardens have succulent collections of internationalimportance.

Kirstenbosch National Botanical Garden TheKirstenbosch National Botanical Garden, located on theslopes of Table Mountain near Cape Town, was foundedin 1913. The garden occupies 528 hectares (1305 acres), ofwhich some 36 hectares (89 acres) is cultivated, while theremainder is maintained as a natural flora reserve.Approximately 6000 indigenous species are grown in thegarden, of which 1500 are succulents. There are a fewindividual plants of three CITES Appendix I species:Aloe pillansii, A. polyphylla, and A. thorncroftii.

47

Approximately 95 per cent of all the succulent plantsgrown at Kirstenbosch are documented in that all theiraccession details and location within the garden are fullycomputerised. Voucher specimens of a number of theplants in cultivation are placed in the ComptonHerbarium, which is also at Kirstenbosch. The systemcould use some improvements on informationaccessibility.

Part of the succulent collection is grown outdoors intwo main areas, namely the Mathews Rockery and theMesem Banks (covering an area of approximately 3000m2), while the remainder, especially the smaller speciesand the research collections, are grown in glasshouses inthe nursery area. The winters in Cape Town are far toowet for many of the succulent species to be grownoutdoors. As a result, a display glasshouse is being built(covering 1400 m2), the main feature of which will be thesucculent flora of southern Africa.

Many of the plants grown at Kirstenbosch are theresult of staff collecting activities, but other collectorswho have made significant contributions to the collectioninclude M. B. Bayer, H. Hall, C. A. Smith, F. Stayner, andW. Wisura.

The Karoo National Botanical Garden, at the foot of theBrandwacht Mountains near Worcester, was founded in1921 but transferred to its present location in 1945. Thisgarden concentrates on plants from the arid semi-desertareas of southern Africa, particularly the succulents.Approximately 11 hectares (27 acres) of the garden havebeen developed for the cultivation of plants, and theremaining 143 hectares (353 acres) are being retained as aflora reserve, protecting a typical example of the localKaroo vegetation, which includes over 80 species ofsucculents.

The lower part of the garden is devoted to summerrainfall arid-area plants, while in the upper part of thegarden their winter rainfall counterparts are grown. Thereare plantings of related species, such as the aloes andmesembs, and elsewhere there are plantings that reflectthe floras of different regions, such as Namibia, theRichtersveld, the Knersvlakte, Tanqua Karoo, LittleKaroo, etc. In all, some 6400 plant species have beenestablished, of which over 2500 are succulents.

There are a number of important referencecollections, e.g. the Huworthia collection, the Conophytumcollection, the Euphorbia collection, and the collection ofsucculent asclepiads, which are grown under controlledconditions in shade and glasshouses. Over 300 species ofrare or endangered succulents are cultivated in thegarden, including two CITES Appendix I species: AZoepillansii (40 plants) and Pachypodium baronii (only 1 plantand which comes from Madagascar). Approximately 80per cent of the collections are fully documented. A smallreference herbarium is maintained at the garden, butmost of the specimen vouchers are sent to the ComptonHerbarium at Kirstenbosch.

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Accession data for all the species in cultivation areavailable. Unfortunately, the plant records system has notbeen kept fully up-to-date and many plant labels havebeen lost or misplaced, making it difficult to relate someplants to their accession data. Attempts are being made torectify this situation. Specimen vouchers of many of theplants in cultivation in the garden are placed in theNational Herbarium located within the garden. This is thelargest herbarium in Africa, with over 1.3 millionspecimens; all the specimen label information is fullycomputerised. The Herbarium also includes the MaryGunn Library, which has an excellent collection of workson succulent plants.

There are a large number of threatened plants incultivation at this garden, including 20 CITES Appendix Ispecies: 15 species of Aloe, 2 species of MadagascanEuphorbia, and 3 species of Madagascan Pachypodium. Inmost cases there are only one or two plants in cultivation,but for some species there are large numbers; forexample, there are 10 plants of Aloe descoingsii, 18 of A.par&a, 12 of A. uauhii, and 55 of A. suzannae.

The major collectors who have made significantcontributions to the Pretoria collection include D. Cole,R. A. Dyer, D. S. Hardy, D. de Kock, J. Lavranos, G.Prinsloo, H. Toelken, and J. van Zanten.

8. Sttidtische S u k k u l e n t e n - S a m m l u n g ZiirichMythenquai 88, CH-8002 Zurich, Switzerland. Telephone:41-l-201-45-54; FAX: 41-l-201-55-40.

The City of Zurich Succulent Plant Collection wasfounded in 1931 and is operated by the city. It consists ofan area of 0.5 hectare (1.24 acres), with nearly all thecollection being housed in glasshouses. The succulentcollection numbers some 26,000 accessions, amounting tomore than 50,000 individual plants. Approximately 20 percent of the collection is from horticultural origin with noindication of locality. The other 80 per cent have somedocumentation; 45 per cent of all accessions are directlyfrom the wild (many in the form of seeds); 10 per cent arelater propagations of unknown status (mostly supplied byhorticultural firms with a collection number); theremaining material is either from seed obtained throughcontrolled pollination, or propagated from cuttings.

Although it has not been calculated, most (if not all)CITES Appendix I succulents are in cultivation. Thecollection also has 14,000 herbarium specimens and aCactaceae seed collection, all of which are documented.

The Zurich Succulent Plant Collection is the officialrepository of the IOS, with its archives, library holdings,and voucher specimens of several research projects.

Collectors who have contributed substantially to theZurich holdings include E. F. Anderson, P. Bally, U.Eggli, J. Lavranos, W. Rauh , W. Rausch, W .Reppenhagen, F. Ritter, and D. Supthut. In addition,numerous accessions have been received from theInternational Succulent Institute (ISI; see under TheHuntington), as well as obtained or purchased from

horticultural firms such as Kohres, Uhlig, Abbey Garden,and from private individuals.

Botanic garden librariesMost of these botanic gardens have important succulentlibraries, which also play a significant role in plantconservation. Plant sciences libraries and librarians inbotanic gardens provide important data for plantconservation both through their own individual plantliterature collections, as well as through the links of theirlibraries with records in other botany library collections.Bibliographies and records on both the literature ofbotany and seed catalogue collections are two importantholdings in libraries. Two such works which pertain torare plant conservation published by The Council onBotanical and Horticultural Libraries (CBHL) are:

1)

2)

Nursery and Seed Catalogs: A Directory of’ Collectiorzs(in North America). 1985. Compiled by June Rogierand Mary Lou Wolfe, Librarians, AndersenHorticultural Library (Minnesota) and PennsylvaniaHorticultural Society (Pennsylvania) in co-operationwith the National Agricultural Library (USA) andAgriculture Canada (Canada).Endangered Plant Species of the World and TheirEndangered Habitats: A Compilation of the Literature.1985. By Meryl A. Miasek and Charles R. Long,Library Director, Indiana University (South Bend)and Director of the Library, The New York BotanicalGarden (New York).

As an example of the type of information available inbotanic garden libraries, the Desert Botanical Garden inthe USA maintains a collection in the Richter Library ofall seed exchange programs in which they haveparticipated since they began a seed exchange programmein 1965. That is also the first year of publication of theDesert Botanical Garden’s Zndex Senzimm. Also, its livingplant and herbarium accession records are maintained inthe Richter Library, dating back to the incorporation ofthe Garden in the 1930s. Most of these are now in acomputer database.

Many botanic gardens also publish records of theircollections, which are useful in ex situ research. Plantnames, locations, old and new maps of plant discoveries,and other records are important information for c’x situresearch. The inclusion of botanic garden libraries andlibrarians in ex situ conservation activities helps insurethat these records are preserved for future use.

Hobbyist collections and nurseries

Whereas traditionally one thinks only of botanic gardensand arboreta when considering conservation collections,private hobbyists and commercial nurseries also provideimportant ex situ collections of succulents. The combinedactivities of succulent plant growers and botanic gardens

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with significant collections of succulents, in large partthrough the encouragement of the InternationalOrganization for Succulent Plant Study (IOS), are nowresolving some of the impediments listed by Given (1987),namely inadequate documentation and conflicting aims.Moreover, co-operative efforts among institutions andindividuals, where they do occur, are making these plantsavailable for research and re-introduction.

Several hobbyists, who are not professional botanists,have significant collections of certain groups ofsucculents. These collections have extensivedocumentation and have been used in several cases formajor taxonomic studies. The 10s has recognised theimportance of these private collections, and hasestablished a Section titled 10s Generic ReserveCollections. For many years the collections at Zurich,Switzerland and Linz, Austria have been recognised assuch, and in 1978 four other gardens received thisrecognition. The 10s set up qualifications and objectivesto be met by 10s Generic Reserve Collections (Taylorand Hunt 1988): “a collection must contain material ofsubstantial scientific importance, especially with regard tosystematic research and/or conservation in their widestsenses.” The objectives are (Taylor and Hunt 1988):

1) Emphasis on material of known wild origin; allaccessions to be properly documented, with details ofdonor, field data, and recipients of propagations;secure labelling of plants and regular updating ofrecords is essential.

2) Preservation of voucher material (dried or liquid) forstudy by taxonomists working with the groupconcerned, and a photographic record of specimensprior to preservation, or as an alternative topreservation. If the Reserve Collection lacks its ownherbarium facilities, preserved materials should bedeposited in a well known and responsible institutionand a record kept of what has been deposited.

3) Representation of as comprehensive a range ofspecies as possible and willingness to assist specialistswho are members of 10s in their research by way ofaccess or loan/gifts of material as appropriate.

4) Active propagation and distribution of rare,scientifically valuable, and endangered species.

5) Willingness to co-operate in studies of floral biologyand vegetative growth under artificial conditionstowards the objective of making rare or endangeredspecies available to a wider public and therebyrelieving the pressure on wild populations.

The following private collections have been approved orare under consideration by the 10s as Generic ReserveCollections (Taylor 1991a):

l Andrea Cattabriga, Bologna, Italy - dwarf MexicanCactaceae

l Keith Grantham, Luton, UK - E@zovhia

50

0

0

l

0

0

0

0

0

0

l--

Ben Groen, Wageningen, The Netherlands -Conophytum, AstrolobaAlan Hart, Cheshire, United Kingdom - CeropegiaFred Kattermann, Sussex, New Jersey, USA -Eriosyce, CopiapoaMassimo Meregalli, Torino, Italy - CopiapoaRoy Mottram, Thirsk, UK - CZeistocactus, OreocereusDavid Parker, Birmingham, UK - EchinocereusHans Till, Attersee, Austria - GymnocalyciumBernd Ullrich, Pohlheim, Germany - AgaveRichard and Franziska Wolf, Wienderwald, Austria -Mammillaria

rred Kattermann and his collection of Eriosyce is anexample of a dedicated hobbyist performing exhaustivefield work, making detailed records, and writing anoteworthy monograph of the genus (Kattermann 1994).Kattermann spent over 45 weeks in the field during 1977-94, legally collecting approximately 1500 plants fromabout 500 different populations. Living material wasdistributed to several institutions for propagation, andherbarium specimens were prepared and given to theDesert Botanical Garden and The New York BotanicalGarden. In addition, he collected living plants of 20 othergenera, most of which have been given to the DesertBotanical Garden. He has willingly provided material toresearchers throughout the world.

Astrophytum asferias in a cactus nursery, Kurashiki,Japan.

Other collections have been critical for taxonomicstudies. The Meregalli collection was the basis for anextensive paper on Copiapon (Meregalli 1991) and theEchtnocereus collection of David Parker was used by NigelTaylor to supplement his field work in his significantmonograph of the genus (Taylor 1985).

Clearly, hobbyist collections are a significant source ofgermplasm of rare and endangered succulents. Theprogramme established by the 10s to insure the propermaintenance and documentation of these collections mustbe encouraged, as well as the implementation of aneffective networking among researchers, institutions, andthe hobbyists.

Some commercial nurseries specialise in thepropagation and sale of specific groups of succulents.Many of their propagated plants are derived either fromseed or cuttings, from field collected, or documentedmaterial. These stock plants, and succeeding generations,can provide a significant source of germplasm for rare andendangered succulents. However, while sometimesreducing demand on wild stock, nursery collections do notnecessarily contribute to the genetic and demographicmanagement of threatened plants. Nursery collections ofpropagules and their artificial propagation for commercialpurposes must go hand in hand with providing safeguards,through botanic gardens and seed banks, for tracking andinsuring that propagules will be available for research andlong-term preservation, and thus contributing to ex situconservation.

Several commercial nurseries have stocks of raresucculents with documentation. Mesa Garden in Belen,New Mexico provides an example of the roles that someof these commercial businesses have in both in situ and exsitu conservation. This nursery arose from Steven Brack’sstrong interest in growing succulents from seeds, and wasstimulated by his membership in the African SucculentPlant Society and its biennial seed distribution. In 1973the nursery began using seeds collected from habitatprimarily within the state and documenting their sourcefor sales. Starting in 1975, he began to make extensivecollections of seeds in Mexico. These collections form thecore of Mesa Garden, and their first seed list of 1980 hadapproximately 1000 offerings.

In 1980 Brack joined with Steven Hammer, a specialiston Old World succulents, to collect seed, especially of themesembs, in South Africa. They encountered the veryrare Conophytum angelicae on one of their trips and wereable to collect nine seeds. From those few propagulesMesa Garden has now produced several generations ofthe plant, with a production of several thousand seedsannually. This has significantly reduced the impact ofcollectors decimating the remaining in situ population ofthis species. Brack has also assisted the US Fish and

Guimar nursery, Tenerife, Canary Islands.

Wildlife Service in the propagation of a rare cactusPediocactus knowltonii for re-introduction back into itsnatural habitat.

From its very beginning, the Mesa Garden nursery hasemphasised the propagation and sale of seedlings andseeds with habitat data. In several instances seed of thesame species are offered from different localities, thusproviding germplasm varia bility. Mesa Garden now has acompreh ensive co1 lecti on of mes embs, as well ashundreds of cacti from both North and South America.This nursery has provided research materials to numerousscientists throughout the world.

It should also be noted thaactivities of botanic gardens and nurseries havethat several succulent species, now virtually extinc

t the cooperative Ed situmeant

t in thewild because of extensive habitat destruction, are widelyrepresented in cultivation. Two examples are Astl’)yhytumasterias and Echinocactus grrasonii.

The author would like to acknowledge the following people whoreviewed and contributed information to this section: SalvadorArias M., Steven Brack, Jane Cole, Dr Urs Eggli, Holly Forbes, DrCraig Hilton-Taylor, Fred Kattermann, Mike Maunder, PeggyOlwell, Liz Slausen, Jean-Marie Solichon, John N. Trager, DrPeter S. Wyse Jackson.

51

Chapter 3

Regional Accounts

Tropical AfricaSara Oldfield

The vegetation of Africa includes sixteen broadvegetation types recognised by White (1983). Woodlandand grassland are predominant. Succulent plants arefound in most vegetation types but are rare in the highforests of West Africa and uncharacteristic of miombowoodland.

White (1983) recognises ten regional centres ofendemism for Africa. Succulent plants are particularlyimportant in the Somalia-Masai, Cape, Karoo-Namib,East Malagasy, and West Malagasy regional centres of

endemism. The Somalia-Masai region, covering Ethiopia,Somalia, Kenya, and northern Tanzania is the main areaof succulent plant diversity in tropical mainland Africa.Between this region of East Africa and the rich southernAfrica flora there is a marked decrease in succulent plantdiversity determined primarily by climatic factors. Incentral Africa there is a much more marked summerrainfall allowing the build up of woody biomass and a longenough dry season to allow regular burning of the groundlayer. As a consequence countries such as Tanzania,Zambia, and Zaire are relatively poor in succulents. Inthis region the Asclepiadaceae, for example, are poorlyrepresented.

Box. 3.1 Main African vegetation types and their succulent diversity

WoodlandOpen stands of trees at least 8 m tall, with a canopy cover of 40 percent or more, with a field layer usually dominated bygrasses. Nearly all types are deciduous or semi-deciduous with some evergreen species. Woodlands are widespread intropical Africa and are especially characteristic of the Sudanian and Zambezian regions with their continental climatesand moderate precipitation falling in the summer. Succulents are relatively poorly represented.

Transitional Scrub forestIntermediate between forest and bushland and thicket and often dominated by tree-like species of Aloe and Euphorbia.

Bushland and thicketBushland includes open stands of bushes, usually between 3 and 7m tall and with a canopy cover of 40 percent or more;in thicket the stands of bushes are closed. Both types are found under a wide range of climatic and edaphic conditionswhich are unfavourable for tree growth. They are most frequent in areas where annual rainfall is 250-500mm and ofirregular occurrence or where there are two pronounced dry seasons. Deciduous bushland and thicket is extensivelydeveloped in the Somalia-Masai region; evergreen and semi-evergreen bushland and thicket is found in the Cape,coastal east and south-east Africa and associated with the drier types of montane forest.

ShrublandOpen or closed stands of shrubs up to 2m tall. The most extensive shrublands are in the Karoo-Namib region and in theCape (fynbos). Shrublands also occur in montane and Afroalpine regions. In the latter they are typically dwarf and formbut one component of a diverse range of communities.

Grassland’ Land covered with grasses and other herbs, either without woody plants or the latter not covering more than IO percentof the ground. Edaphic grasslands are widespread throughout Africa and include vast areas in the Serengeti (developedon volcanic deposits and maintained as grassland by grazing), grasslands associated with seasonally or permanentlywaterlogged soils and also secondary grassland, which has replaced forest or woodland after human intervention (suchas burning and cultivation).

Wooded grasslandLand covered with grasses and other herbs, with woody plants covering between IO and 40 percent of the ground. Thisis the most widespread vegetation in the Sahel and in the Kalahari part of the Kalahari-Highveld zone. It is also commonin the Sudanian and the Zambezian regions.

Deserts and semi-desertsArid landscapes with a sparse plant cover, except in depressions where water accumulates. Semi-desert vegetationbegins to occur when the mean annual rainfall drops below c. 250mm, e.g. in parts of the Karoo-Namib, Somalia-Masai,and the Sahel and on the margins of the Sahara Desert. True deserts include the Sahara, the floristically richer NamibDesert, and parts of northern Kenya.

52

In recent years tropical rain forest vegetation hasreceived much conservation attention. Arid vegetationtypes, however, may also be very rich in plant species andunder threat. The Karoo, for example, an area of semi-desert, contains a wealth of succulents and otherxerophytic plants. Similarly, arid areas on theSomalia/Kenya/Ethiopia border and in the rainshadows ofthe Eastern Arc mountains in Tanzania (see Lovett 1988)exhibit high levels of endemism at the level of the plantspecies. Fynbos, the evergreen bushland and thicket of theCape region, has the highest concentration of species perunit area in the world: 8550 vascular plant species in89,000 km2 (Goldblatt 1978).

The vegetation of Africa has been modified by humaninfluence over many thousands of years. In recent yearsthe scale of destruction of semi-arid, woodland, and forestecosystems has assumed major proportions. Thedeterioration of dry lands may not perhaps be soimmediately obvious as forest destruction, but it isnevertheless as serious, both for local inhabitants and formaintenance of plant species diversity. Generally, the bestpreserved semi-arid and arid vegetation types can now befound only in remote areas or in national parks, far fromany artificial water supplies.

Somalia-Masai regional centre of endemism

The Somalia-Masai vegetation group of White (1983)includes virtually all Somalia, much of Kenya andEthiopia, together with the arid northern portion ofTanzania and north-western Uganda. Within this sectionthe succulent floras of Ethiopia, Kenya, and Somalia areconsidered further. The entire Somalia-Masai vegetationgroup is considered to be one of Africa’s ten regionalcentres of endemism. There are about 4000 vascular plantspecies in the region with 31 per cent endemism. Thereare many endemic succulents (e.g. stapeliads).

The main vegetation type is deciduous bushland,dominated by Acacia and Commiphora. This can varyfrom open bushland 3-4 m tall to almost impenetrablethicket some 10 m tall. A subtype on limestone andgypsum includes many local endemics. Enclaves withinthis type are riparian forest and semi-evergreen bushlandon hills and the lower slopes of mountains. Semi-desertgrassland and shrubland occur in low rainfall areas.

Overgrazing caused by large-scale herding of animals(cattle, camels, goats) has transformed the originalvegetation in many areas. This has become particularlysevere in recent years due to increase in population andrecurrent droughts. Areas near permanent wells orwaterholes are the most affected. Also, over vast areas,the deciduous bushland has been converted to secondarybushed grassland by temporary agriculture or the cuttingof fuelwood for charcoal-burning.

Less than five per cent of the unit is included inprotected areas (MacKinnon and MacKinnon 1986).Many large national parks have been established ineastern Africa, mainly for the protection of spectacular

concentrations of wildlife in savanna habitats. There are,however, no conservation areas in the Ogaden region ofEthiopia, Kenya, and Somalia, for example, which arcimportant for succulents.

Kenya

Kenya has in total about 6000 flowering plant species.Inventories of species are relatively complete with themajor exception of the undercollected north-east region.The full distribution of most plant species is, however, notknown. Three-quarters of Kenya’s land area comprisesarid or semi-arid ecosystems. Plant endemism is high,especially if regional endemics are included in the totals(National Biodiversity Unit 1992). Inselbergs in semi-aridareas are botanically important sites. There are 364indigenous taxa of succulents in 22 plant families reportedfor Kenya (Wabuyele, in Zitt. 1997; see Box 3.2).

Threats to the succulent floraHabitat destruction through processes such as agriculturaldevelopment, building construction, and expansion is themain threat to succulent plants in Kenya (Newton 199s).Grazing is a particular threat to succulents such asCaralluma spp. which are not protected by spines or thepresence of unpalatable substances such as the latex ofEuphorbia spp.

Commercial exploitation is another significant factoraffecting certain succulent species. Collection of Kenyansucculents for the export market is not thought to takeplace on a major scale, but there is some collection of rareand unusual succulents for specialist growers overseas.Undescribed species of Raphionacme are, for example,currently offered by a specialist succulent trader inGermany. Other species of Asclepiadaceae have alsobeen threatened by overcollection including Huernia spp.,Caralluma spp., Echidnopsis dammanniana, andEdithcoZea grandis.

Exploitation of native Aloe spp. has also been a threatto wild populations. Plantations of AZoe spp. were firstestablished near Mombasa in 1986 following thePresidential decree protecting wild populations fromharvesting for leaf exudates. Unregulated initial stockingand restocking of these plantations is probably stilldepleting wild Aloe populations. A plantation in northernKenya was established by transplanting wild plantsresulting in considerable harm to wild populations(Newton 1991). Field observation has shown that the lawis rarely observed and there is evidence of continuingillegal harvesting. In some areas of Kenya harvesting fromthe wild appears to do little harm to populations becausealmost all defoliated plants survive. In other areas, it hasbeen reported that wild plants have been completelydestroyed by harvesting activity. In the Baring0 area ofKenya, collection ofAloe leaf is causing serious damage towild populations. Local people are paid Ksh 20 for 20litres of leaf extract (fl = Ksh 66) which would involveharvesting several hundred plants (Newton, in Zitt. to S.

53

Oldfield, WCMC, 1992). The main Aloe species exploitedin Kenya is A. secundifloru, although A. turkanensis is alsothought to be used. Exploitation is all for export, with nocompany using Aloe exudate to manufacture productswithin the country (Newton, in Zitt. to S. Oldfield, WCMC,1992).

The conservation status of Kenyan succulentsVarious partial lists of threatened plants have beenproduced for Kenya. The most recent list compiled withinthe country includes succulent species in the familiesCrassulaceae, Aizoaceae, Portulacaceae, Euphorbiaceae,Asclepiadaceae, and Liliaceae (Luke 1991). WCMC holdsrecords of 82 threatened succulent species of Kenya in thelatter three families. Recent consultation with theNational Museums of Kenya Plant Conservation andPropagation Unit resulted in a target list of species knownto be under threat in their natural habitat, though theconservation status of many others are not known(Newton 1995). In this list 67 taxa have been given toppriority status, in most cases because of threats to habitats(Annex 4). Data sheets on three Kenyan succulents areincluded in the IUCN Plant Red Data Book (Lucas andSynge 1978). These are Caralluma distincta, C. tubiformis,and Euphorbia wakefieldii. These remain top priorities forconservation attention.

Priority sites for conservationUpland grassland areas, especially those with impededdrainage ‘vleis’, support interesting and very rare plantspecies such as Brachystelma keniense and should beprotected.

Existing conservation measuresKenya has a protected area network covering over sevenper cent of the total land area. The protected areas arebiased towards savanna/semi-arid areas with significantpopulations of large mammals. Most protected areassuffer damage through encroachment, poaching,pollution, or overuse by tourists (National BiodiversityUnit 1992). Desert areas are inadequately represented inthe protected area system. Small areas of lava desert areprotected in the Marsabit and S. Turkana National Parks.More areas for protection should be identified in theChalbi and Koroli deserts (Mackinnon and Mackinnon1986).

There are various categories of protected areas inKenya. Under the Wildlife (Conservation andAmendment) Act 1976, as amended, the following may bedesignated:

0 Nationaliniurv. 0

Parks - vegetation is protected from cutting,r setting fire: no clearance or cultivation is

aliowJed.” /

Box 3.2 Kenya Succulent Species Conservation Project

This Project, initiated mid-1996 under the direction of the Plant Conservation Programme (PCP) of the East African Herbarium,holds as its objective to facilitate conservation of Kenya’s succulent plant taxa. The working document for this project is a reportby Prof. Len Newton (1995) entitled “Succulent Species in Kenya”, prepared as a contribution to this Action Plan. In the reportNewton documents 364 succulent taxa in 22 families in Kenya. Here 67 taxa, representing 12 families, are listed as being rareand /or endangered (Annex 4). These taxa comprise the target PCP list for formulation of conservation action. There are 23 taxafor which the conservation status is unknown and these will receive primary attention.

The project objectives, some tasks for which have already been completed, are as follows:l Document type specimens and duplicates for succulent plant species in the East African Herbarium,l Determine how many type localities have been destroyed and whether those remaining are threatened,l Develop a specimen database for threatened taxa,l Assess the pressure on wild populations and the present conservation status of taxa known from only one locality,l Collect germplasm for ex siitu conservation and propagate in cultivation those taxa that are rare and under threat in their

natural habitat, andl Document species in trade and / or cultivation as a guide to monitoring exploitation pressure.

Field surveys are aimed at verification of the extant populations of targeted taxa and evaluation of their current conservationstatus; documentation of threats to each of them; collection of stocks for ex situ conservation and collection of herbariumspecimens for further research.

It has been noted from analysing the collection database that there is a lack of adequate and accurate information ondistribution of most taxa. Collections occur mostly around Central Kenya, the lower Rift Valley, and Western Kenya. In order toreach any meaningful consensus on the conservation status of these taxa it will be important to take complete inventory of theunder-collected regions and protected areas. Areas of high succulent diversity should be identified and subsequentlyconserved.

In relation to the trade of succulents, the Project wishes to develop a dataset of species in cultivation in nurseries, conductsurvey on international traded for the Kenyan taxa, and develop a list of rare and endangered taxa for Cl1-ES in country policing.

a

54

National Reserves - less strictly controlled with otherland uses permitted, for example, traditional grazingrights and water rights are protected.Local Sanctuary - vegetation is not protected.Protection area - adjacent to national parks, nationalreserves, or local sanctuaries, they are legally declaredto protect species, habitats, and ecology.

Under the Forests Act 1942, as amended, Forest Areas orCentral forests are designated and also Nature Reserveswhich are strictly protected sites within forest reserves.

Succulent plants are represented within many ofKenya’s protected areas. Usually, however, plant speciesinventories are not available, so it is difficult to assess towhat extent rare and threatened species are protected.

Examples of protected areas important for succulentplant conservation include the following:

”

Mount Kulal Biosphere Reserve - succulent EuphorbiaSPP*Amboseli National Park - four types of semi-aridvegetation.Hell’s Gate National Park - situated south of LakeNaivasha in the Rift Valley. There is a wide variety ofsucculents in the area.Mount Suswa - proposed extension to Hell’s GateNational Park, in Nalvasha District.Mutomo Plant Sanctuary - degraded site covering 16ha in Kitul District. This was first established in 1964as a joint enterprise between the County Council ofKitul and the Kenya Horticultural Society. TheSanctuary protects a range of interesting succulentplant species in situ and also provides a site for ex situconservation of plants from other semi-arid parts ofKenya.

Ethiopia

The flora of Ethiopia consists of about 5765 floweringplant species of which lo-20 per cent are endemic.Endemism is particularly high in the sub-desert Ogaden inthe south-east as well as in the forests of the south-west.There are also local centres of endemism in themountains of Ethiopia.

Ethiopia shows a considerable variety of habitats, mostof which have at least some succulents growing in them.In very simplistic terms the country can be divided intofour main areas:

Central highlands - the Semien Mountains rise to 4600m in the north and in the south there is an extensiveplateau at over 4000 m. Convenient divisions can be madeat very approximately 1500 m as the bottom limit, and atabout 3500 m between a lower Afromontane area and anupper Afroalpine area.

The Afroalpine areas are occupied by Erica bushlandat lower altitudes and moorland at higher altitudes and in

areas of poorer drainage. Succulents are restricted to afew species of Aloe, all endemic, and a variety ofCrassulaceae, with several endemic species of Sedlrpn,Umbilicus botryoides, the very distinctive endemicRosularia simense, Aeonium leucoblepharum and theendemic genus Hypagophytum semiense. The latter twoalso extend down into the Afromontane zone whereconditions permit.

The Afromontane area is mostly densely populatedand intensively cultivated with only remnants of theoriginal vegetation below 3500 m. Many of the lower partswere probably at one time forested, but now onlyremnants of the original forest are left. These wereprobably mostly dominated by Juniperus procera and 01~61europea ssp. afiicana. In the south and west the rainfall ishigher and more reliable and Podocarpus gracilior is oftena major component of the forests whilst in the extremesouth-west, at rather lower altitudes and higher rainfall, adepauperate Guinean - Congolan forest type is still to befound characterised by various species of Celtis andTrilepisium madagascariensis. The forested areas arerelatively poor in succulents though the consistentlyepiphytic endemic Sedum epidendrum is a notableexception and Euphorbia ampliphylla can be an importantcomponent tree, reaching a height of at least 24 m.Euphorbia ampliphylla is often used to form living hedges.Succulents are most common on rocky slopes and it seemsrather clear that some groups, most notably various Aloespecies have benefited from the creation of such habitatsby forest clearance. Kalanchoe, including the endemics K.schimperiana and K. petitiana, are often common atthicket margins.

Western lowlands - The flora of these areas has verystrong affinities with the flora of West Africa with manyspecies reaching their easternmost limit in Ethiopia. Thearea is not well known, but the indications are thatrelatively few succulents occur. Much of the area isoccupied by wooded grassland subject to regular burningwhich will kill the majority of succulent plants. Portdacafoliosa s. str. is one succulent that does occur and providesa good example of the western affinities of the area - thetype was collected in Ghana. The name has been used inEast Africa, but closer examination has shown that suchmaterial is not conspecific with the West African plant.

Eastern lowlands - The area between the highlands andthe Red Sea is mostly occupied by the Afar Depression, atrue desert with vast salt deposits and rather few plants ofany kind. Ecologically, the most important group ofsucculents are members of the Chenopodiaceae which areoften dominant in the moister, usually saline areas. Somestapeliads are found in suitable situations, but these arenearly all species much better represented to the south,only Caralluma edulis has not been recorded fromelsewhere. Similarly, a few species of Euphorbia are foundbut only E. triaculeata is restricted to the area. There is atendency forthe eastern escarpment of the highlands to

55

receive a significant portion of their rainfall in winter and succulents is deciding on what should be included, asthis is reflected in the occurrence of plants with distinct there are large numbers of semi-succulent species. A listMediterranean affinities. Amongst these is an ephemeral of some important genera within the families containingSedum related to S. hispanicum. succulents is presented in Table 3.1.

Southern and south-eastern lowlands - These include awide variety of deciduous bushland and woodlandbelonging to the Somali-Masai vegetation group. The vastmajority of Ethiopian succulents are found in this area,with only the Crassulaceae and possibly Aloe betterrepresented elsewhere in the country. The dominant treesand shrubs throughout the region are species of Acaciaand Commiphora, but species of succulent Euphorbia areoften dominant locally. There are many areas which arerich in succulents and even areas where stapeliads such asCaralluma penicillata are locally subdominant.

The succulent floraThe major problem in drawing up a list of Ethiopian

Threats to the succulent floraCommercial exploitation - To date there is relatively littleevidence of specific commercial exploitation ofsucculents. The notable exception may be Euphorbiaabyssinica which in the past was the primary source ofmatchwood in Ethiopia. The current situation is unclearbut there are still large populations of this species over amajor part of the country. The commercial collection ofAloe for the pharmaceutical industry appears to have beenvery restricted, in marked contrast to Kenya.

The biggest potential threat must be from succulentcollectors. Some Ethiopian endemic succulents areamongst the most desirable of all collectors’ plants, mostnotably Euphorbia piscidermis and E. gymrzocalycioides,

Table 3.1 Succulent flora of Ethiopia

Aizoaceae Seven genera and perhaps 13 species, only 3 or 4 species of Delosperma are of horticultural interest.Only 1 of these, D. nakurense, is relatively well known.

Aloaceae 38 species of Aloe, 22 endemic, mostly at higher altitudes, some known from rather few localities, butnone vulnerable and some extremely abundant.

Asclepiadaceae

Cactaceae

Many non-succulent.

Rhipsalis baccifera native to south-west forests; Opuntia at least 3 species growing as weeds, oneexploited locally for fruits.

Crassulaceae

Cucurbitaceae

Ethiopia has the greatest number and diversity of Sedum spp. of any African country (Gilbert 1985).

Most species not succulent; Momordica rostrata and Cephalopentandra ecirrhosa with well-developedsucculent caudexes, and Mukia pallidinervia a true leaf succulent.

Euphorbiaceae Three genera include succulents: Euphorbia, Monadenium, and Jatropha. Euphorbia is the largestgenus within Ethiopia, and also includes the largest number of succulents. The exact number dependslargely on what definition is used for succulence, there being a number of marginal species. The largestgroup are the 45 species, 15 endemic, belonging to the all succulent subgenus &phobia; subgenusLacanthis has 4 species, 3 endemic; subgenus Esula has 3 good succulents plus a number of marginalsucculents,

Geraniaceae Most species of Pelargonium at least slightly succulent; only one of the Ethiopian species, P.boranense, a local endemic from southern Ethiopia, of interest to general collectors because of thesucculent stem and very large bright red flowers.

Passifloraceae Two species of Adenia having true succulents - A. venenata and A. aculeata; others subsucculent, mostnotably A. ellenbeckii, or not succulent.

Piperaceae

Portulacaceae

Eight species of epiphytes, none endemic, all widespread in Africa and sometimes further afield.

Talinum: 3 species, 7. porfulacifolium common and widespread; Calyptrotheca somalense at leastlocally common in western Sidamo and Gamo Gofa - apparently with a short flowering period andprobably under-recorded; Portulaca: 15 species, some rather poorly known, others widespread,including a troublesome pantropical weed.

Vitaceae Most non-succulent, no succulents endemic, some species of Cissus leaf-succulents (C. rotundifolia) orstem succulents (C. Cactiforme, C. quadrangulare, C. quinquangulare); Cyphostemma betiforme astem succulent restricted to gypsum areas of the south-east.

56

and there is no doubt that most these would be veryvulnerable to any kind of commercial collecting. It is vitalthat such exploitation should be very strictly controlled.

Agriculture - Habitat degradation from agriculture isobviously a threat to many succulents, but it is probablynot of very great importance within Ethiopia. Mostspecies are found in areas not suitable for cultivation andindeed some species, most especially some of the moreshrubby Aloe species, have probably extended their rangesin parts of the country where denudation by erosionfollowing cultivation has created extensive open rockyslopes.

Overgrazing - Overgrazing is a problem in some areas,but in many cases succulent species survive surprisinglywell wherever there is some protection by rocks andunpalatable shrubs.

Introduced species - Opuntia cf. dillenii and two or threeother species are a major weed problem in some areas butthere is some evidence to suggest that native succulentsare at least sometimes able to benefit from the shelterthese spiny plants can offer.

Conservation status of succulentsA comprehensive listing of the conservation status ofEthiopian succulent plants has not yet been prepared.WCMC holds records of 76 nationally threatenedsucculent species of Ethiopia.

Existing conservation measuresMost major conservation areas have been identified andprovisional or nominal protection implemented.Ethiopia’s protected areas consist of national parks,sanctuaries, wildlife reserves, and controlled huntingareas. Only two national parks, Awash and SimenMountains, have been legally gazetted.

All the national parks contain some succulents.Political upheavals and lack of resources have made theeffectiveness of these parks open to question, but there isno doubt that they will offer some protection fromagricultural development, probably the greatest overallthreat to succulents within Ethiopia. The SemienMountain National Park includes the type localities ofmany Ethiopian high-altitude endemics includingRosularia simensis, Hypagophytum semiense, and probablyAloe steudneri. Awash National Park includes the typelocalities of Pachycymbium sacculata and Euphorbiaawashensis. Quite a number of other succulents also occurwithin this park.

The Mago National Park, which is not yet gazetted,has Euphorbia grandicornis, E. scoparia, Sansevieria spp,and Adenium obesum. Nechisar National Park, also notyet gazetted, has fine stands of Euphorbia tirucalli.

Somalia

Somalia has approximately 3000 flowering plant species ofwhich about 500 are endemic. Highly specialisedvegetation types within the country support manyendemic xerophytic plant species suggesting that aridclimatic conditions have remained unchanged for longperiods of time.

The vegetation of Somalia consists mainly of Acacia-Commiphora deciduous bushland and thicket particularlyin the south. There are large areas of semi-desertgrassland and deciduous shrubland in the north andextending south along the coast.

The north-east of Somalia (Cal Madow mountains) isparticularly rich in succulent plant diversity and isconsidered to be an internationally important centre ofplant biodiversity (WWF and IUCN 1994). Special&dlimestone habitats, with outcrops of pure gypsum,harbour many endemic species. Some of these specieshave been known only from single collections by earlynaturalist explorers and many others have only beendiscovered in the past fifteen years. The bush-coveredplains of southern Somalia have succulents in commonwith south-east Ethiopia and the better-known region ofnorth-east Kenya. There are also local endemics confinedto isolated coastal outcrops or coastal dunes.

Threats to the succulent floraDesertification threatens species of the succulent scrub.Somalia has the greatest proportion of pastoralists inAfrica, and livestock accounts for 40 percent of thecountry’s GNP. Overgrazing is the dominant threat tonatural habitats and the succulent species which theycontain.

Some succulent species are able to withstand grazingpressures. Aloe spp. are, for example, generallyunpalatable. Aloe megalacantha invaded extensive areasfollowing removal of woody vegetation by livestock, forexample, on the plains surrounding the war-stricken cityof Hargeisa. Other species of AZoe, such as A. pir-ottae andA. peckii are, however, adversely affected by grazingpressures. These two species only grow in the shelter ofbushes and once the vegetation is removed the speciesbecome progressively rarer. Other succulent species areparticularly susceptible to grazing pressures, for example,Asclepiads which are not spiny or protected by poisonoussubstances. Euphorbia columnaris, endemic to a smallarea in north-east Somalia is almost extinct becauseovergrazing has led to soil erosion. Plants of the specieshave weak root systems which are unable to supportmature plants once the soil is eroded.

War in the Horn of Africa has lead to the disruptionof traditional grazing regimes leading to furtherenvironmental instability. The flow of refugees intonorthern Somalia has lead to destruction of vegetation forkilometres around the camps (Hutchison 1991).

57

Table 3.2 Succulent genera of Somalia

Family Genus No. ofspecies

Aizoaceae (Aizoon) 1Sesuvium 2Trian thema 5[11

Aloaceae Aloe 27Apocynaceae Adenium 3Asclepiadaceae” Cal0 tropis 1

Caralluma 17+2 (7endemic)

Ceropegia 6Cryp tolepis 3+1Curroria 2Cynanchum 8Diplos tigma 1Dregea 3Duvalia 3Echidnopsis 12+1Edithcolea 1Glossonema 4Gomphocarpus 3Huernia 3Kanahia 1Leptadenia 3Odontan thera 1Orbea 1Oxys telma 1Pen tarrhinum 2Pen ta tropis 2Pergularia 2Periploca 3Pleuros telma 1Pseudolithos 4Pseudopectinaria 1Raphionacme 1Rhytidocaulon 2+1Sarcostemma 2Secamone I+1Socotora 1Strobopetalum 1Tacazzea 1Tenaris 2Vince toxium 1Whitesloanea 1

Crassulaceae Aeon&m 1Cotyledon 1Crassula 3Kalanchoe 12Sempetvivum 1Umbilicus 1

Euphorbiaceae Euphorbia 99+3[5](>50%endemic)

Monadenium 3Portulacaceae Anacampseros +I

Calyp tro theta IPortulaca WITalinum 3+1

Note: * in need offurthertaxonomic work

Source: Kuchar I985

Conservation status of succulentsAlthough a comprehensive listing of the conservationstatus of succulent plants of Somalia has not yet beenprepared, IUCN categories have been applied to anumber of succulent species, especially in the generaEuphorbia and Aloe. WCMC holds records of 61nationally threatened succulent species for the country.Data sheets on Euphovbia cameronii and Whitesloaneacrassa are included in the IUCN Plant Red Data Book(Lucas and Synge 1978). The genus Pseudolithos isendemic to Somalia. All four species of this genus are rareand severely threatened. In general, many of theasclepiads of Somalia are considered to be exceedinglyrare (&char 1986). The low representation of stapeliadsin herbarium collections may be because the smallpopulations initially located have been decimated byintensive grazing and erosion (Kuchar 1986).

Priority sites for succulent plant conservationFour centres of endemism and plant diversity for Somaliaare given in Centres of Plant Divers@ (WWF and IUCN1994). These are Hobyo, the Cal Madow mountain range,the fixed dune vegetation in southern Somalia, and theNugaal Valley.

Hobyo - The area around Hobyo (Obbia) in the MudugRegion of central Somalia, covering around 3000 km’,consists of a coastal plain with dunes and areas oflimestone pavement, particularly in the north; a lowlimestone escarpment running more or less parallel to thecoastline; and a low plateau further inland. The limestoneescarpment reaches about 440 m south-west of Hobyo. Itis dissected by deep gorges which run approximately eastto west. The gorges contain seasonal streams. Thevegetation of the coastal plain is open and treeless with alow vegetation of grasses, herbs, and shrublets. Thelimestone escarpment is covered with a species-richAcacia-Commiphora bushland and the inland plateau alsohas bushland and woodland dominated by species ofAcacia and Commiphora. The area is still relativelyinaccessible and in need of further botanical study.Succulents are frequent. The threats to the flora are notparticularly severe at present, although the effects ofover-grazing and cutting of woody vegetation forfuelwood are damaging in some places. Fortunately, themost botanically interesting vegetation, that of the dunesand limestone gorges, is still largely intact. Parts of thearea have been proposed as a Game Reserve.

Cal Madow - As mentioned above, the Cal Madow (AlMedu) mountain range, a large area which extends rightup to the north-east corner of Somalia, is aninternationally important centre of plant endemism. Thisremote area is relatively sparsely populated and wasapparently untouched by the recent war. The regionconsists of a coastal plain bordering the Gulf of Aden,locally known as the “Guban”, and an uplifted limestoneplateau lying to the south and dipping to the south-east.

58

The plateau scarp reaches 2416 m at Shimbiris (thehighest point in Somalia), to the north-west ofCeerigaabo.

The vegetation of Cal Madow varies greatly accordingto altitude and rainfall and includes coastal plain of desertor semi-desert, with little or no vegetation; the subcoastalzone with sparse to dense, primarily woody vegetation;slopes of the escarpment covered by a macchia-likeevergreen or semi-evergreen scrub which, at higherelevations, grades into remnants of Juniperus forestdegraded by logging. To the south of the escarpment,vegetation consists of severely degraded open woodlands.

Examples of phytogeographically interesting succulentspecies present in Cal Madow include A e o n i u mleucoblepharum and Euphorbia balsamifera. Severalsucculent Euphorbia spp. are endemic.

The most interesting vegetation along the escarpmentis still largely intact and threats to the flora are notparticularly severe at present. Daalo Forest Reserve, tothe north of Ceerigabo, and the surrounding mountainarea have been proposed for protection as a nationalpark. However, at present there are no functioning naturereserves in Somalia.

Fixed dune vegetation in southern Somalia - Thisvegetation supports xerophytes and succulent species ofmany genera. Large, fixed and vegetated dunes occur insouthern Somalia along the Indian Ocean from south ofHobyo to just south of the border with Kenya. These aremostly lo-15 km wide and between 20 and 60 m high.Some dunes have been deprived of their vegetation byovergrazing and are moving inland. The vegetationconsists of Acacia bushland. There are more than 200vascular plant species, with over 20 strict endemics and anumber of endemic subspecies. Some of this vegetationshould be protected by the proposed Lag BadanaBushbush National Park.

Nugaal Valley - An arid area along the Wadi Nugaal,about 250 km long. The western part consists of massivedeposits of gypsum and anhydrites, while the coastal partis mainly limestone. The vegetation consists of open semi-desert grassland or bushland. The succulent flora isparticularly rich with a number of local endemics. Prioritysites associated with the Wadi Nugaal include the gypsumhills around Las Anod where the sparse degradedvegetation supports a rich endemic succulent flora withAloe inermis, Dorstenia gypsophila, Adenia aculea ta,Raphanocarpus stefaninii, Euphorbia columnaris,Pterodiscus, and Caralluma spp. The Las Anod NationalPark has been proposed by Bally and Melville (1973) toprotect fine scenery, rich and varied flora, and the Somaliwild ass (Equus asinus somalicus). They also suggest that afenced floral reserve be created for Euphorbia columnaris.The limestone plateau to the north of Eil is another arearich in succulent plants including many endemics and hasbeen proposed as a national park by Bally and Melville(1973).

Other prioritv succulent sites include:Gaan Libah Forest Reserve - Aloe jucunda, Aloehildebrandtii, Euphorbia abyssinica, Kalanchoe “pp.,Echidnopsis spp.; proposed as a national park by Ballyand Melville (1973).Sheikh Pass area in the Golis Mountains - rich inendemic species; Whitesloanea crassa, Pseudolithoscubiformis, Euphorbia phillipsiae, Aloe somaliensis,Aloe hemmingii, Euphorbia inculta, Edithcolea grandis,M o n a d e n i u m ellenbeckii, Kleinia gunnisii, andEchidnopsis cilia ta.Molidera Hills - two gypsum hills 58-62 km south ofErigavo on the road to El Dab; Aloe mohderana,Pelargonium christophoranum, Dorstenia gysophila, andDracaena ombet; floral reserve suggested by Bally andMelville (1973).Bulo Burti - limestone ridge with rich flora includingMonadenium stellatum; floral reserve suggested byBally and Melville (1973).

Existing conservation measuresSomalia lacks any organised protected area system toconserve its critical sites. Of the conservation areasestablished and proposed since 1969 only Balcad andAlifuuto Nature Reserves are currently functional, andplans have been prepared for the proposed Lag Badana-Bushbush National Park. The legal establishment of allconservation areas in Somalia would result in more thaneight per cent of the total area being covered. In addition,there are 27 ‘protected forests’ and grazing reservesestablished to protect grazing lands (WCMC 199 1).

The author would like to thank Mike Gilbert and Susan Carter-Holmes for the provision of information and suggestions for thisaccount; the Plant Conservation Programme of the NationalMuseums of Kenya has also provided valuable information. TheProtected Areas Unit of WCMC is also thanked for the provision ofinformation.

MadagascarSara Oldfield and Diedrich Supthut

Madagascar has an extraordinary flora, rich in endemictaxa. Out of an approximate total of 7600 endemic plantspecies, over 600 are succulent plants. The succulent floraof the island is of great value botanically, ecologically, andeconomically, and yet it is one of the most endangeredsucculent floras of the world.

The succulent plants of Madagascar are listed inAnnex 7. The family Didiereaceae (see Box 3.3) isendemic to Madagascar and a number of succulentgenera in other families are restricted to the island. Newsucculent species and varieties continue to be discoveredand, despite the level of international botanical andhorticultural interest in the group, the taxonomic status ofmany Madagascan succulents remains poorly known.

59

Box 3.3 Didiereaceae, a succulent family endemic to Madagascar

The Didiereaceae Drake is a small family endemic to Madagascar consisting of four genera, Alluaudia, Alluaudiopsis, Decay/a,and Didierea, with eleven species. The spiny cylindrical or conical pachycaul stems of the plants give a superficial resemblanceto columnar species of Euphorbia or cacti and the Didiereaceae are sometimes known as “the cacti of the old world”. The stemsare woody, and scarcely succulent, with limited space for water storage. The plants have simple, deciduous leaves, except forAhaudia dumosa, and small, unisexual flowers. Accounts of the family are given by Rauh (1963) and Rowley (1993).

Species of the Didiereaceae are ecologically important components of the dry thorny forest in the south and south-west ofMadagascar. Much of this vegetation has been cleared for agriculture and most of the remaining stands are impenetrable ornearly so. Some remnant patches of dry thorny forest occur within protected areas. The Reserve naturelle integrale deTsimanampetsotsa, for example, has vegetation dominated by Didiereaceae and Euphorbiaceae and covers part of the veryrestricted distribution of Ahaudia montagnacii. The Reserve naturelle integrale d’Andohahela has a good representation ofAlluaudia and Didierea spp.

The whole family is threatened mainly by habitat destruction and utilisation of the wood for construction and charcoal production.The wood of Alluaudia procera, A. ascendens, and A. montagnacii is used locally in the areas where the species grow and is soldin the cities of Madagascar. Exploitation of the wood is a threat, for example, to the species and vegetation in the Reservenaturelle integrale d’Andohahela.

Another threat to Didiereaceae is collection for horticulture. All species are in demand by collectors, but they rarely flower incultivation, and seed is in short supply. Some species are cultivated in Madagascar for export, but others such as Didiereamadagascariensis and D. trolh’i are gathered from the wild and exported as seedlings. The whole family has been included inAppendix II of CITES since the Convention came into force in 1976. This has enabled the collection of trade data for the family,which revealed, for example, the import of thousands of wild plants of Didierea and Alluaudia into Europe in the mid-1980serroneously labelled as artificially propagated plants. ,

Flare de Mudaguscar et des Comores (Humbert 1936) of Madagascar, although a general identification guide tocovers most of the major succulent families, but many ofthe early volumes are out of date. The genus Euphorbiahas not yet been covered by the Flora. Some of therelatively well known succulent groups such as the genusAloe and geophytic Euphorhia spp. are in need oftaxonomic attention (Supthut and von Arx 1993). In thecase of Aloe spp. their spatial separation on isolatedinselbergs, long intervals of time between speciesdescription, and difficulty of access to sites have allcontributed to taxonomic uncertainty. Professor Rauh hasprepared the first volume of Succulent and XerophyticPlants oj’ Madagascar (1995) and volume two is expectedin 1997. There is currently no local guide to the succulents

the common plants of the island is being prepared.Madagascar can be divided into a number of

phytogeographical regions reflecting local differences intopography, geology, soils, and climatic conditions.Within four broad regions - eastern, central highlands,western, and southern / south-western - seventeenvegetation types are recognised, within which there aremany distinct plant communities. No systematicclassification at the community level is available as yet.Most of Madagascar’s vegetation has been extensivelymodified and only about 20 per cent of the naturalvegetation remains.

lnselberg on the CentralPlateau, Madagascar.

60

The eastern escarpment to the central highlands allthe way to the coast, characterised by high rainfall and avariety of soil types, is the richest botanically. Vegetationconsists of a band of evergreen rain forest below 800 mand along the coast, now extensively deforested andreplaced by ‘savoka’, or secondary lowland forest.

The undulating central plateau region, from 500-1500m in elevation, is characterised by lateritic clay soils andmontane and cloud forests. The plains have beenovergrazed and burnt for centuries resulting inimpoverished grasslands with their upper layers stronglyeroded and are now very poor in succulent species (Rauh1983b). Rupicolous vegetation occurs on the inselbergswhich occur in the plateau area of central Madagascar.The inselberg rock formations of volcanic origin representlocal ecological niches in the middle of a montane forestzone. The initial colonists, after lichens and mosses, areplants of the genus Fimbristyhs which produce a pureblack neutral to acidic humus carpet. This becomescompletely desiccated in the dry season (Rauh 1983b).Succulents can tolerate the soil conditions and are wellrepresented in the rupicolous vegetation with, forexample, species of Pachypodium, Aloe, Kalanchoe,Cynanchum, Euphorbia, and Tetradenia. Most of thehighland Aloe species are inhabitants of the inselberg rockformations: A. haworthioides; A. parvulu, which grows inthe quartz Itremo rocks in cushions of Fimbristylis or incracks in the rocks; A. Zaeta in the Massif du Mont Ibity

Didierea trolli, Madagascar.

growing in association with A. ibitensis; A. parallclifolia;and A. trachyticolu (Rauh 1983b). The floral compositionof rupicolous vegetation on the western rocky outcrops is,in general, relatively poorly known.

The western region has climax vegetation of drydeciduous forest with woody endemic succulents ofgenera such as Adenia, Adansonia, Uncar-ina,Pachypodium, and Euphorbia. Most of the deciduousforest has been destroyed by human activity and isreplaced bY secondary or wooded grassland. The westernescarpment of the central plateau consists of lowevergreen sclerophyllous forest, and transitional forest inthe Sambirano region of north-west Madagascar.

The southern and south-western region 1s

characterised by arid climatic conditions. The climate ischaracterised by lack of seasonality with the slightprecipitation (approximately 300 mm per year) fallingirregularly throughout the year. Towards the coast highair humidity is a climatic feature, with heavy dew and fogprecipitation. The vegetation consists of deciduousxerophytic thicket, sometimes known as spiny desert. Thisvegetation is dominated by Didiereaceae and Euphorbiaspp. It has a sparsely developed ground layer consistingpredominantly of succulents. Deciduous thicket has beenreplaced by grassland over much of its natural area ofdistribution.

Succulent plants are represented in allPhYtoge% raphical regions of Madagascar. The two mostimportan t regions are the Central Plateau, on theinselbergs, and the dry forests in the north-west, west, andsouth . The remarkable vegetation of the southern coastalregion supports the highest percentage of endemicsucculent plant species for the country as a whole.

Threats

Burning - Large areas of Madagascar have been burnedsince the first settlers arrived on the island 1500 years ago.The extensive areas of grassland are burned each year toprovide pasture for zebu cattle. Some succulent speciessuch as Aloe macroclada and various species of Euphorbiasuch as E. primulifolia are relatively fire-resistant, and theinselberg communities initially escape damage from fire,but in general burning is a major threat to the succulentplant flora. Pachypodium spp. are for example particularlysusceptible to fire damage as are some rare Aloe spp.

Other Madagascan succulents considered to beendangered by fire include the asclepiads: Ceropegiadimorpha, C. armandii, both of which are also sought afterby succulent collectors, C. bosseri, C. Zeroyi, andCynanchum rossii. The latter two have not been seen inthe wild for the past twenty years. The genus Stapelianthuswhich is confined to the south-west of the island, is alsosusceptible to fire, with the following species endangered

s by burning and clearance of the xerophytic forest:E5 Stapelianthus decaryi, S. pilosus, S. madagascariensis, S.I/)i montagnacii, S. ins&is, S. hardyi, S. keraudrenae, S.d arenarius, and S. calcarophilus.

Grazing - The zebu cattle of Madagascar and theirimpact on the flora and fauna are well known. There arean estimated 10 million zebu on the island and the cattleare of great cultural importance.

Clearance for agriculture - Traditional agriculture inMadagascar consists of various forms of shifting andsettled cultivation producing predominantly rice, withcassava, sweet potatoes, and maize. The main cash cropsare coffee, grown mainly along the east coast and in thenorth-west; cocoa, also grown in the north-west; vanilla,grown mainly in the north; and cloves grown on theeastern coastal plains. Large plantations with oil palms,green peppers, cinnamon, and cloves are found in thearea.

Agricultural development has been less important inthe arid succulent-rich areas of southern Madagascar.Nevertheless, a large area of Didiereaceae vegetation hasbeen cleared in the Amboasary region for sisal, Agavesisalana, plantations. Production of sisal declined untilrecently but is now expanding again and remains animportant industry. Two new large sisal plantations havebeen laid out in the vicinity of Amboasary and anothernew sisal plantation exists north of Morandava. The rarespecies, Euphorbia cylindrifolia, E. ambovombense, E.ampanihense, together with some Stapelianthus and smallAloe species grow in Alluaudia-Euphorbia forests close tosisal plantations and could be destroyed throughintensified agricultural development together withextensive clearance of forests for charcoal production.Opuntia plantations are also established in this region.

Introduced species - Agave sisalana is an invasiveintroduced species which continues to have a detrimentalimpact on the ecology of the xerophytic thicket

Euphorbia quartziticola, a Rare plant restricted toquartz substrates; threatened by habitat destruction.

vegetation. The species spreads by prolific vegetativereproduction, destroying primary vegetation.

The prickly pear, Opunfia dilenii, first introduced toFort-Dauphin as fencing around houses in around 1770,quickly became established to the detriment of indigenousvegetation. Brought under control in the 1920s thespecies is again established in southern Madagascar.Various other Opuntia spp. are planted for fodderthroughout southern Madagascar.

Furcraea is another invasi ve, and increasinglycommon, introduced species, particularly on the CentralHigh Plateau.

Alluaudia-Euphorbiaforest, since destroyed,featuring E. plagiantha,A. ascendens, andA. procera.

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Alluaudia montagnacii, used for construction andcharcoal production.

Local use - Collection of fuelwood affects woodysucculent species in the vicinity of towns such as Tulear,Sakahara, and Fort-Dauphin. The production of charcoalalso consumes significant quantities of woody succulentsparticularly close to Tulear, where there is a commercialcharcoal operation. Charcoal production is also commonto the north of Tulear and at Ampanihy, Tsihombe,Amboasary, and Ambovombe. Species used for charcoalproduction include Euphorbia spp., Alluaudia spp., andDecaryia madagascariensis. Charcoal production is bothfor domestic consumption and export overseas.

Woody succulent species are utilised as timber forconstruction of dwellings. This places a strain on wildpopulations of various species, notably AlZuaudiaascendens and A. montagnacii. Another species ofDidiereaceae, Decaryia madagascariensis, is heavilyexploited by local people. Its trunk forms dense, firmsections of charcoal which are highly valued, and thethinner twigs are used for hedging and corral building.Species of Didiereaceae are also used as live fencing.

A considerable number of succulent species areutilised locally for medicinal purposes as documented, forexample, by Jenkins (1987). Certain species are rare, butit is not known to what extent collection for medicinal usehas a detrimental impact on wild populations. Some

succulent species are also considered to be sacred in someparts of Madagascar; for example, Pachypodiumbrevicauze is avoided by local inhabitants.

Collecting for horticulture - Madagascan succulents arc ofhorticultural interest to specialist collectors worldwide.The international demand for succulent species fromMadagascar has led to the wholesale removal of wildplants for export primarily to Europe, Japan, and theUSA. Around 100 succulent species have been exportedon a fairly regular basis, the principal genera beingPachypodium, Aloe, and Euphorbia. It is clear that wherethere is a monetary value to plants as a result ofinternational horticultural demand, there is a strongincentive to collect and export the material which is notseen as having any intrinsic value locally. Both bulkremoval of common species for the ‘supermarket’ tradeand collection of very rare species for the specialistmarket continue to take place. The latter is probablymore threatening or damaging to local populations.

The actual species collected for export changeaccording to fashions in succulent trade. At present thereis a strong demand for caudiciform plants, most of whichare not listed on the Appendices of CITES. At the sametime newly described species of all genera will be subjectto particular threat.

Conservation status

There has been no systematic attempt to categorise theconservation status of Madagascan succulent species.IUCN categories have been applied to various succulentgroups during the 1980s based on the field knowledge ofindividual experts, and this information was recorded inthe Plants Database now maintained by WCMC.Conservation categories are given for Madagascansucculents in Annex 7 where these are known. For themajority of species there is limited distribution data andeven less site assessment information on which to basecurrent conservation status. Taxonomic uncertainties addto the difficulties of applying conservation categories.

The main stimulus for field-based assessment of the

Pachypodium rosulatum var. stenanthu sold along atourist route.

63

conservation status of succulent species known to be rarehas been the listing of certain species on Appendix I ofCITES. During 1992 and 1993 two field surveys werecarried out in Madagascar by D. Supthut and B. von Arxunder the auspices of CITES (Project No. S52). The fieldwork involved training of local conservation personnel insucculent species identification. Three sets of herbariumspecimens were made for deposition at the DEFherbarium in Antananarivo; Stadtische SukkulentenSammlung, Zurich, and Conservatoire et Jardinbotanique in Geneva. Further survey work of this natureis a priority for the conservation of the Madagascansucculent flora.

Priority sites for conservation

Madagascar’s succulent species occur throughout theisland in all major vegetation types. There are certainareas which are outstanding in terms of their succulentrichness and diversity:

1) The whole area of southern xerophytic thicketcommunity, dominated by Didiereaceae and shrubbyor arborescent Euphorbia spp., and resticted to a smallcoastal region about 50 km wide, is an internationalconservation priority. According to Rauh (1978d),“All in all, the vegetation of the precipitation-poorsouth-west Madagascar is because of its numerousmorphological, biological, floristic specialities and itswealth of endemism, something so unique in the worldthat all effort should be made not only to guard theseplants but to preserve them for posterity.” Not manyuntouched, coherent areas remain in the south andsouth-west, but within the area certain sites can beidentified as particularly outstanding for succulentconservation. One good area is between Ampanihyand Ejeda (along Route Nationale 10) down to thecoast at Itampolo and Androka.

2) Another important area of the southern xerophyticthicket community is that to the south of Tulear where

Saint Augustin, south-west coast, an importantsucculent habitat featuring Uncarina, Moringa,Euphorbia, and Stapelianthus.

3)

4)

5)

6)

7)

In 1992, this small population of Aloe calcairophilawas totally removed by collectors.

the succulent flora is particularly diverse. One siteidentified as a priority for conservation is to the east ofthe coast road, from the Tulear exit to the junctionwith the road to the interior, towards Saint Augustin.Rare species of Aloe, Euphorbia, Stapelianthus,Kalanchoe, Moringa, and Operculicarya occur.

A third important area for this type of vegetation isSite d’interet biologique du Lac Anony. Aloe suzannaeand A. helenae grow in this area, together withEuphorbia cylindrifolia and E. fkancoisii at the westernend of their distributions. Succulent species ofCucurbitaceae such as the heavily traded Xerosicyospubescens also occur.

Site d’interet biologique de Zombitse has a tropicaldry climate with 750 mm annual precipitation. Theforest is heavily exploited.

Mount Ibity in the Central Plateau is a very importantsite for succulent plant conservation with endemicssuch as Aloe trachyticola, Pachypodium brevicaule, andEuphorbia primulifolia. It is regularly visited bysucculent plant collectors.

Co1 d’Itremo, another site in the Central Plateau,hosts an exceptionally rich flora and fauna due to itsdistinctive geology and soils. Euphorbia quartzicola,Pachypodium brevicaule, P. densiflorum, Aloecalcairophila, and Aloe compressa occur there.

Site d’interet biologique du Nord de Toliara PK 32contains primary forest to the north of Mora-Mora -spiny forest with Didiereaceae, Pachypodium, andarborescent Euphorbia spp. Unfortunately, large areasof forest were cleared in Autumn 1994.

64

Adansonia fony,Didiereamadagascariensis,Euphorbia stenoclada,and De/onyx floribunda;type locality ofAlluaudiopsismamierana.

8) La Montagne des Francais is important forpopulations of Adanso~iu suarezensis, an endemic tothe north of Madagascar, which is threatened byhabitat destruction at its other locations. The site alsohas Pachypodium baronii var. windsori, P. decalyi, P.ruthenbergianum, Euphorbia viguieri, E. neohumbertii,Aloe suarezensis, and Impa tiens tuberosum.

Existing conservation measures

The conservation of Madagascar’s biodiversity is widelyrecognised as being of outstanding importanceparticularly by international conservation agencies. Majorprogrammes of conservation activities such as theMadagascar National Environmental Action Plan(NEAP) have, therefore, been developed, largely withoutside funding. There has been, in general, remarkablylittle attention paid to the conservation needs ofMadagascar’s rich succulent flora.

Protected areasProtected areas currently cover approximately 10,350 km’(1.76 per cent) of Madagascar’s land area and Malagasybiodiversity is not fully represented. Of particular concernfor succulent plant conservation is the gap in protectedarea coverage in the south-west. In total only around50,000 ha of spiny forest vegetation is protected (Pollock1986). Also of major concern is the fact that none of thesucculent-rich inselbergs of the Central Plateau areprotected. In general, the protected areas which do existon pape r are not adequately prot.ectedMost do not have botanical species inven

on thtories,

e ground.and these

are urgently required.Five different categories of protected area are

65

recognised:

a>94d)e>

Reserves naturelles integrales (Strict NatureReserves),Parts nationaux (National Parks),Reserves speciales (Special Reserves),Forets classees (Classified Forests),Perimetres de reboisement et de restauration(Reafforestation Zones).

The establishment of Strict Nature Reserves dates back to1927. The relevant legislation is Decree 66-242. Underthis legislation access to the sites is strictly forbiddenother than for scientific research purposes. National Parklegislation is contained in Decrees 58-07 and 62-371.Access to National Parks is controlled and exploitation ofsome forest products by local people is allowed. SpecialReserves have been set up by a series of decrees and aredesigned to protect certain plant or animal species. Access toreserves is free but some damaging activities, including thecollection of natural products, are forbidden.

A review of ecosystems, protected areas, and speciesconservation requirements carried out by WWF andMadagascan conservation authorities has recommendedthe expansion of the protected area system (Nicoll andLangrand 1989). The lack of adequate protected areacoverage in the south-west is noted but no specific sitesfor conservation are identified. Out of 16 critical siteslisted as meriting protection, only one in the north ofMadagascar is an important succulent area: La Montagnedes FranCais, an area of dry limestone forest withdifferent vegetation to the nearby Montagne d’AmbreNational Park. It is also planned to give the Site d’interetbiologique de Zombitse and the Vohibasia forest, thestatus of Reserves speciales. These sites are transitionalbetween western and southern vegetation and areremnants of the species rich Foret de Sakaraha which hasvirtually disappeared.

Protectedconservat1) Part national de 1’Isalo - encompasses part of the

rusandston e Isa lo Massif,abundant with Puchypodiu

picolousrosula turnm

vegevar.

tation isgracilius,

2)

3)

4)

5)

Euphorbia primulifolia var. begardii, Ceropegiadimorpha, E. isaloensis, and Aloe isaloensis. The areaneeds to be extended to include land south of theroute national No. 7 and north of the Onilahy River.Reserve naturelle integrale de L’Andringitra - rich

on theendemic flora with Aloe and Kalanchoe spp.crags. A species list does not exist for this area.Reserve naturelle integrale de 1’Ankarafantsika -Pachypodium rosulatum var. drakei, P. sofiense,Euphorbia cremersii, E. pauliani, E. pedilanthoides, and

*ale du Tsingy de Namorokad crassulacean plants withPachypodium ambongense.

Reserve naturelle integrale de Tsimanampetsotsa -

areasion

important for succulent

vegetation dominated by Didiereaceae, for example,Decaryia madagascariensis, Alluaudia procera, A.montagnacii, A. humbertii, tree Euphorbiu spp.,Pachypodium spp., and Adansonia fony.Reserve naturelle integrale d’Andohahela - has agood representation of Alluaudia and Didierea spp.with Didierea trollii, A. procera, A. ascendens, A.humbertii, and tree Euphorbia spp.Reserve naturelle integrale du Tsingy de Bemaraha -Euphorbia moratii (3 vars.), many Aloe spp. ,Lomatophyllum spp. and Pachypodium spp.Reserve speciale Cap Sainte Marie - the limestonerocks support Alluaudia comosa, Alluaudiopsisfiherensis, Aloe millotii, Euphorbia capsaintemariensisand Crassula humbertii.Reserve speciale Bezaha-Mahafaly - a 500 ha reserveprotecting xerophytic vegetation in south-westMadagascar, with Didiereaceae, Aloe, Euphorbia andPachypodium spp.

10) Reserve speciale d’Ankarana - Pachypodium decutyi,Euphorbia ankarensis, E. herman-schwartzii, E. alfiedii,E. pachypodioides, E. neohumbertii, and Cyphostemmalaza.

Many of Madagascar’s protected areas have receivedmore intensive biological research than other parts of thecountry, but the more inaccessible sites are still poorlyexplored. The preparation of detailed inventories of theprotected areas would help in the analysis of currentprotection given to threatened succulent species inMadagascar. Renewed inventory work is currently beingcarried out in actual or proposed protected areas, mainlyunder the auspices of Missouri Botanical Garden’sMadagascar Research and Conservation Program. This,however, concentrates on wet forests with little work onthe dry or high plateau flora.

National legislationThere is currently no legislation giving special protectionto rare and threatened succulents in Madagascar.Protected area legislation has been discussed above.Conservation legislation relating to vegetation and flora

66

Box 3.4 Conservation agencies and botanical institutions in Madagascar

Direction des Eaux et For&s (DEF): This government department, under the Ministere de la production animale (elevage etp&he) et des Eaux et for&, has responsibility for forest management and protected areas. DEF is also the CITES ManagementAuthority.

Office National de L’Environnement (ONE): ONE is a newcross-sectoral review of policies which relate to the environment.

Government body, created in 1991 I It is currently planning a

Agence Nationale pour la Gestion des Aires ProtkgBes (ANGAP): ANGAP is an NGO, established with funding fromUSAID, which works closely with DEF. ANGAP has taken responsibility for 34 out of 50 areas designated for protection and isinvestigating the development of management plans for protected areas.

National Association for Environmental Actions (ANAE): ANAE is a recently established NGO which serves to providefinancing to communities, local NGOs and village organisations for watershed management, reforestation, and erosion controlactivities developed through local initiatives,

WWF-Madagascar: WWF representation in Madagascar was established in 1979 under a legal decree. Since that time WWFhas implemented activities relating to education and public awareness and more recently implementation of field activitiesparticularly relating to management of protected areas. Botanical work at present relates mainly to ethnobotany.

Jardin Botanique de la DRST Tsimbazaza: maintains important living collections of Madagascan plants, but does not havesuitable climatic conditions for the cultivation of many of Madagascar’s endangered succulents. There is a growing core of welltrained and experienced botanical staff, supported by a strong reference library and herbarium. It has been suggested that aCITES Scientific Authority should be based at the Garden.

in general consists of: a law of 15 January 1930 protectingforests; an ordinance, No. 75.014-1975, bringing CITESinto force. Law No. 91.008 of 25 July 1991, concerninganimal protection, reinforces the importance of CITESbanning the export of animal species listed in theAppendices, but does not refer to plant species.

Knowledge of the status in the wild of succulent plantspecies is essential in determining which plants should belisted for legal protection. At present this information isonly partially available.

Review and additional information for this section were kindlyprovided by Blaise Du Puy, Martin Jenkins, and Bertrand vonArx.

Southern AfricaCraig Hilton-Taylor

Southern Africa (including Botswana, Lesotho, Namibia,South Africa, Swaziland, and Zimbabwe) has long beenrecognised as an area of remarkable plant diversity withextremely high levels of endemism (see Cowling et al.1989; Cowling and Hilton-Taylor 1994; Gibbs Russell1985; Goldblatt 1978; Good 1974). The high levels ofdiversity and endemism found in the flora in general, alsoextend to the succulent plant groups as has been shown byBarkhuizen (1978) Court (1981) Van Jaarsveld (1987)and Smith et al. (1993) ( see Table 3.3). Considering thatsouthern Africa is a predominantly warm-temperate,semi-arid region with an overall mean annual rainfall ofless than 400 mm, it is not surprising that there are a largenumber of succulent plants in the flora. What is perhaps

most astonishing, is the extent to which the majorsucculent families have speciated in southern Africa,particularly the Aizoaceae, Aloaceae, Asclepiadaceae,Crassulaceae, and to a lesser extent, the Euphorbiaceae(see Table 3.4). This diversity can in part be attributed tocontemporary ecological conditions, particularly thesubcontinent’s transitional location relative to thesubtropical summer rainfall and winter rainfall climaticzones, combined with a complex topography andheterogeneity of geology and soils which allow for steepecological gradients (Cowling and Hilton-Taylor 1994). Inaddition the region’s history of aridity (see Van Jaarsveld1987) combined with a special genetic potentiality(Ihlenfeldt 1994) has also undoubtedly played a majorrole in promoting speciation in the succulent groups. Theheterogeneity of southern Africa’s flora is well illustratedby its classification into five phytochoria (Werger 1978;White 1983) seven biomes (Rutherford and Westfall1986) and, in South Africa alone, 70 major vegetationcategories or Veld Types (Acocks 1953).

Succulent plants have long been utilised by people insouthern Africa; however, it was not until the start of theactive botanical exploration of the region by the earlyseventeenth century explorers and naturalists, that thissucculent wealth came to the attention of people inEurope and eventually the rest of the world (Smith et d.1993). Despite the rises and falls in the popularity ofsouthern African succulent plant species amongsthorticulturists, botanists and hobbyists around the world,the demand for these plants has undoubtedly had anegative impact on the wild populations. This demand,together with the enormous impact of people inmodifying and transforming virtually every ecosystem insouthern Africa (Macdonald 1989) has resulted in a

67

number of southern Africa’s succulent species becomingthreatened with extinction. It is these rare and threatenedspecies which form the focus of this section.

There have been several attempts to describe andcatalogue the succulent flora of southern Africa. Some ofthe earlier descriptive works include those of Rawe (1968)and Barkhuizen (1978) which were followed by the morecomprehensive work of Court (1981). Van Jaarsveld(1987) provided a review of all the succulent plantfamilies in southern Africa including their distribution,reproductive and defence mechanisms, and adaptations todrought. This work was complemented by that of Smith etal. (1993) which focused largely on the taxonomic issuesin southern African succulent families. Other publicationswhich have helped document the southern Africansucculent flora include Jacobsen (1960, 1974) Percy-Lancaster and Richards (1991) and the recentlypublished works of Eggli (1994) and Eggli and Taylor(1994). In addition to the above works there is a largenumber of books, scientific papers, and popular articlesdealing with specific southern African succulent familiesand genera (see Smith et al. 1993 for references to someof these). All of the above publications are helping tobring us closer to compiling a complete ‘register’ of allsouthern African succulents (Smith et al., in press).

A major inconsistency among many of the publicationslisted above is the definition of succulence. Some authorshave followed a fairly broad definition, whereas others arevery strict. The differences in numbers of succulent taxacited in each work can therefore be attributed in part tothe author’s definition. The task of obtaining totalnumbers of succulent taxa is further complicated by thefact that all of the publications dealing with all thesucculents in southern Africa, including the most recent,are taxonomically outdated and in addition there aremany differences of opinion on which taxa are worthrecognizing and consequently which names should beaccepted. For the Flora of southern Africa (FSA) region(the area including Botswana, Lesotho, Namibia, SouthAfrica, and Swaziland), the treatment of Smith et al.(1993) was followed, although it was not recognised thatmany of the numbers of species cited are incorrect, asthey were based on outdated information in Arnold andDe Wet (1993). For Zimbabwe, the checklist compiled byPercy-Lancaster and Richards (1991) was used. Evencomparing Percy-Lancaster and Richards (1991) to Smith

FSA region are presented and discussed separately tothose for Zimbabwe.

Table 3.3 shows the numbers of succulent taxa(families, genera, species) recorded in the FSA region.These numbers were derived from Smith et al. (1993)however, all succulent species which have becomenaturalised in the region (e.g. many species of Cactaceae)were removed from the counts. There are therefore 3377species (in 201 genera and 26 families) of succulent plantsindigenous to the FSA region. (Note: these figures do notinclude the numbers of infraspecific taxa.) If one adds thesemi-succulent species, the total rises to 3873, whichrepresents some 17 per cent of the total flora of the FSAregion (Arnold and De Wet 1993) and 39 per cent of theworld’s 10,000 succulent plant species (Rowley 1978).Table 3.4 gives a breakdown of the major succulentfamilies in the FSA region, and the number of succulentgenera and species in each family in comparison to thenumbers which are non-succulent. From Table 3.4 it isclear that the Aizoaceae is the most important succulentfamily, contributing some 63.4 per cent of all thesucculent species f o u n d i n t h e r e g i o n . T h eAsc lep iadaceae , Aloaceae , Crassu laceae , andEuphorbiaceae all contribute significant numbers ofsucculent species to the flora. Further details on thesefamilies, their genera and species are provided by VanJaarsveld (1987) and Smith et al. (1993).

Table 3.5 indicates that succulents are also animportant component of the flora of Zimbabwe, with 336taxa (species and infraspecific taxa) of succulents,pachycauls, and caudiciforms recorded (cycads andgeophytes listed by Percy-Lancaster and Richards (1991)have been excluded). It is estimated that there areapproximately 6500 species of flowering plants inZimbabwe (Kimberley 1993) therefore some five per centof these can be termed succulent or semi-succulent. Themajor families and genera in Zimbabwe which containsucculent or semi-succulent species are discussed in detailby Percy-Lancaster and Richards (1991) and Kimberley(1993). Comparing the taxa in the list compiled by Percy-Lancaster and Richards (1991) to those listed in Arnoldand De Wet (1993) it is evident that there are 115succulent taxa recorded from Zimbabwe which do notoccur in the FSA region. Therefore the grand total for thewhole of southern Africa (FSA region plus Zimbabwe) is3988 succulent taxa, i.e. nearly 40 per cent of the world’s

et al. (1993) poses problems due to the differing succulents occur in southern Africa.interpretation of succulence. Hence the results for the

Table 3.3 Numbers of succulent taxa (families, genera, and species) in the Flora ofsouthern Africa regionColumn five is the percentage of the 10,000 succulent species in the world. (Source: Smith et al. 1993)

SucculentSemi-succulent

Total

families

2613

39

genera

20155

256

species

3377496

3873

Per cent of speciesFSA world

15 342 5

17 39

68

Table 3.4 A summary of southern African plant families with succulent species, arrangedin decreasing number of succulent species per familyNote that some families are not restricted to southern Africa but, for the purposes of this table, only those species occurring inthe Flora of southern Africa region were taken into account. (Source: Smith eit al. (1993), but some numbers have been updatedby this author.)

Family Number of generanon-succulent succulent

Number of species Percent total Largest succulentnon-succulent succulent succulent genus in the

flora family (No. of species)

Aizoaceae 0 c. 1-UAsclepiadaceae 39 27Aloaceae 0 6Crassulaceae 0 6Euphorbiaceae 47 3Asteraceae 233 IOZygophyllaceae 4 4Asphodelaceae 2 2Portulacaceae 1 5Geraniaceae 3 2Chenopodiaceae 15 3Vitaceae 3 2Apocynaceae 1% 2Pedal iaceae 6 2Passifloraceae 4 1Piperaceae 1 1Dracaenaceae 1 1Lamiaceae 37 1Hyacinthaceae 24 3Dioscoreaceae 0 1Goodeniaceae 0 1Cactaceae 0 1Viscaceae 0 1Brassicaceae 38 1Sterculiaceae 6 1Rubiaceae 61 1

Total 543 c. 205

0 c. 2141 63.4415 c. 264 7.8

0 c. 262 7.80 236 7.0

284 181 5.42232 58 1.7

8 c. 49 1.5107 43 1.3

2 42 1.2c. 263 29 0.9

157 14 0.441 11 0.332 9 0.324 7 0.215 c. 5 0.2

1 4 0.12 4 0.1

233 4 0.1c. 346 4 0.1

11 3 0.10 2 0.10 1 0.03

16 1 0.03158 1 0.03177 1 0.03235 1 0.03

c. 4759 c. 3377 100

Ruschia (c. 350)Stapelia (c. 45)Aloe (149)Crassula (I 5 I)Euphorbia (179)Senecio (30)Zygophyllum (c. 44)Bulbine (4 1)Anacampseros (22)Pelargonium (15)Sarcocornia (9)Cyphostemma (9)Pachypodium (5)Pterodiscus (4)Adenia (c. 5)Peperomia (4)Sansevieria (4)Plec tran thus (4)Bowiea (2)Dioscorea (3)Scaevola (2)Rhipsalis (I)Viscum (I)Heliophila (I)Sterculia (1)Phylohydrax (I)

Distribution in southern AfricaAlthough succulents are found in practically all parts ofsouthern Africa, by far the greatest numbers of species,genera, and families are found in the western arid parts ofthe region (Hilton-Taylor 1994, 1996a; Van Jaarsveld1978) (see also Chapter 1 for details on theirdistributions). In terms of phytochoria, biomes, andvegetation types, succulents within South Africa tend topredominate in the Karoo-Namib Region (see White1983), the Desert, Succulent Karoo, and Nama-KarooBiomes (sensu Rutherford and Westfall 1986) and in thevarious karroid and false-karroid Veld Types (vegetation

Table 3.5 Numbers of succulent, pachycauland caudiciform taxa found in Zimbabwe(Source: Totals modified from Percy-Lancaster andRichards, 1991,)

Families Genera Taxa

Succulent 14 48 231Pachycaukaudiciform 23 8 105

Total 37 56 336

types) described by Acocks (1953). Jurgens (1990) coinedthe term “leaf succulent zone” to describe part of thisarea which is extremely rich in leaf succulent species.High numbers of succulent species are also found in dryhabitats along the east coast of South Africa, extendingfrom the Eastern Cape along the coast of KwaZulu-Natal,through Swaziland into the Eastern and NorthernTransvaal (see Figure 3.1). Succulents generally avoid theinterior plateau above the Great Escarpment because ofthe occurrence of frost (Figure 3.1). Some species,however, are able to survive the harsh highveld winters byestablishing themselves in relatively frost-free niches or bydeveloping a degree of tolerance to frost as for example,in some of the AZoe species (see Hilton-Taylor and Smith1994).

Succulent taxa occur in nearly all parts of Namibiaexcept for the central dune field of the Namib Desert andthe sandy and semi-humid eastern and north-easternparts. Major centres of endemism with concentrations ofsucculent species in Namibia include the Southern NamibDesert (Sperrgebiet or Diamond Area No. 1). theHunsberge, the Warmbad-Pofadder region, theKarasberge, Damaraland, and the Kaokoveld.

In Botswana, although succulents are mostly found in

69

the south-eastern and eastern parts of the country (to theeast of the eastern edge of the Kalahari sands) where theyoccur on suitable rocky outcrops, a number of species,particularly members of the Asclepiadaceae, are able tosurvive in the Kalahari sands area under semi-desertconditions (see Hargreaves 1990 and Plowes 1989).Succulents occur all over Zimbabwe, but the greatestnumbers are in the eastern highlands area (see Percy-Lancaster and Richards 1991).

Conservation status

Data on the conservation status of southern Africansucculent species were obtained from a variety of sources.The primary sources were the Southern African Red DataBook: Plants - Succulent and Nama-Karoo Biomes(Hilton-Taylor, in prep.) and the Red Data List oj’southernAfvican plants (Hilton-Taylor 1996b) and the subsequentupdate (Hilton-Taylor 1996~). The latter is a collaborativeproject with information obtained from the threatenedplants databases maintained by the various conservationagencies in southern Africa (Cape Nature Conservation:R. Stanvliet and T. Anderson; Natal Parks Board: R.Scott-Shaw; Orange Free State Directorate of Nature andEnvironmental Conservation: J. du Preez; TransvaalDirectorate of Nature and Environmental Conservation:W. Boyd; Namibian Ministry of Tourism andEnvironment: M. Strohbach; National Herbarium ofNamibia: G. Maggs; and the Swaziland National TrustCommission: K. Braun). A number of publications on thethreatened plants of specific geographical or politicalregions also provided valuable baseline data, particularlyon the past conservation status of certain taxa. Theseincluded Everard (1988); Fourie (1984, 1986); Hall et al.(1980); Hall and Veldhuis (1985); Hilton-Taylor andSmith (1994); Jankowitz (1975, 1977); Kimberley (1971,1975, 1980, 1991) and Muller (1985). Information was

also obtained from the Plants Database maintained byWCMC, particularly for Zimbabwe. In addition, anumber of amateur and professional botanists wereconsulted and are listed in the acknowledgements.

All the data obtained is stored in a database (known as‘SARARES’) developed at the Ecology and ConservationResearch division of the National Botanical Institute inKirstenbosch, Cape Town. This database is updatedregularly, and it is hoped that it will provide the basis forthe future development of a southern African threatenedplants working group, which will enable better co-ordination of research and conservation activitiesconcerning threatened plants across the subcontinent.Having all the data in one centralised database, alsoenables one to look at the conservation status of eachspecies across its entire distribution range rather than injust certain political or geographical areas. This will helpto prevent limited conservation resources being spentunnecessarily on species which are only locallythreatened. The conservation status as listed in Annex 8,is therefore the global status for each taxon. Only seven ofthe 576 taxa listed are not endemic to the FSA region,and all of these are in effect endemic to the greatersouthern African region in that they occur in theadjoining countries of Angola, Mozambique, and/orZimbabwe, where they are also under threat.

The IUCN Red List Categories (see Annex 16) wereused in describing the conservation status of each taxonconsidered to be rare or threatened. In some cases wherethere was uncertainty as to which category was mostappropriate hybrid categories as used by the WCMC weregiven. A new set of categories with rigorous quantifiablecriteria has been developed (see Mace et al. 1993; Maceand Stuart 1994; IUCN 1994b) but have not been usedhere as much of the southern African data is too sparse toapply the new criteria at this stage. Once consensus hasbeen reached on which taxa should be on the southern

Table 3.6 Numbers of globally threatened succulent taxa in the Flora of southern Africaregion based on information in Annex 8EX = Extinct in the Wild, E = Endangered, V = Vulnerable, R = Rare, I = Indeterminate, K = Insufficiently known,Percent = percentof total number of succulent species in that family in FSA region which are threatened.

Family EX E V R I K Total Percent

AizoaceaeAloaceaeAsclepiadaceaeEuphorbiaceaeCrassulaceaePortulacaceaeAsteraceaeAsphodelaceaeGeraniaceaeVi taceaeApocynaceaePassifloraceaeBrassicaceaeZygophyllaceae

7 14 2720 31

2 3 192 6 172 1 4

21

1 2

9142522839172743

3448231

5089

1313

7

1

1211

Total 14 45 104 289 54 105 611 18%

21

22310593686221IO117423

1

IO4035362650172624362260

1002

70

Figure 3.1. Map of the frequency distribution of succulent plant genera in South Africa, including Lesotho andSwaziland, showing the number of genera recorded per quarter degree grid square. Data extracted from the PRECIS(PREtoria Computerised Information System) database maintained at the National Herbarium (PRE), Pretoria, South Africa.(Map prepared by M. M6ssmer and G.F. Smith).

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African threatened list, all the taxa will be assessed usingthe new criteria and the new categories assigned. Futureadditions to the list will also be evaluated in terms of thenew criteria.

The listings of all succulent taxa (excluding semi-succulents) considered to be rare and threatened in theFSA region and in Zimbabwe are given in Annexes 8 and9 respectively. As the databases on which these annexesare based are constantly changing, the informationpresented for the FSA region was correct as of 31November 1996 and for Zimbabwe as of 19 June 1995.

A summary of the information in Annex 8 is presentedin Table 3.6 (for all taxa with hybrid conservationcategories, the higher status was used). This summaryreveals that 18 per cent (611 taxa) of the succulents in theFSA region are considered to be at risk or threatened. Ofthese, 14 taxa are thought to be Extinct in the Wild and149 are seriously threatened. The overall extinction ratecould in fact be considerably higher as many of the taxaclassified as Indeterminate, may also be Extinct. Most of

the extinctions are directly the result of habitat loss due toagricultural activities, while none can be attributed toremoval by collectors, although many of the taxa in theEndangered and Vulnerable categories are facingextinction as a result of overcollecting. The mostthreatened families include the Aizoaceae (223 taxathreatened), Aloaceae (105 taxa), Asclepiadaceae (93taxa), Euphorbiaceae (68 taxa), and Crassulaceae (62taxa). Although the Aizoaceae shows the greatestextinction rate, it is the Aloaceae which are the mostthreatened with 51 taxa in the Endangered andVulnerable categories. Table 3.7 shows the threatenedstatus of the Aloaceae at generic level. From this it isapparent that although Aloe has the greater number ofthreatened taxa, it is the taxa within Huworthlu which areunder the most threat (see Hilton-Taylor and Smith 1994for details).

Annex 9 shows that 48 succulent taxa are consideredto be threatened in Zimbabwe. Twenty-one of thesethreatened taxa are endemic to Zimbabwe, while eight of

71

Table 3.7 Numbers of threatened alooid taxa in the Flora of southern Africa regionEX = Extinct, E = Endangered, V = Vulnerable, R = Rare, I = Indeterminate, K = Insufficiently known, nt/ne = not threatened or notevaluated, Per cent = percentage of taxa in that genus threatened. (Source: Hilton-Taylor and Smith 1994 and Annex 8.)

Genus

AloeAstrolobaChortolirionGas teriaHaworthiaPoellnitzia

Total

E

4

117

22

V

11

117

29

R

311

541

42

Total ntlne Per cent

54 123 311 4 200 1 09 13 41

40 75 351 0 100

105 216 33%

the non-endemic taxa are also globally threatened, groups by international pharmaceutical companies hasalthough one of these is insufficiently known for complete taken place including succulent taxa like species ofevaluation. The Zimbabwe figures combined with those Commiphora. This exploitation has apparently alreadyfrom the FSA region indicate that 603 of southern lead to the complete destruction of some plantAfrica’s succulent taxa are considered to be globally populations in Damaraland and the Kaokoveld (C. Hines,threatened at present. pers. comm.).

ThreatsBotswanaVery few of the succulents in Botswana occur inconservation areas, and therefore they are all potentiallyunder threat. Fortunately, due to the low populationdensity and relatively undisturbed nature of much of thevegetation, few species are considered to be threatened,although an assessment of the situation is long overdue.Many succulents in Botswana are utilised for a variety ofethnobotanical purposes, e.g. Euphorbia duseimata is usedto induce vomiting after poisoning and in the treatment ofmental disease, stomach pains, and venereal disease;Orbeopsis knobelii is eaten raw or roasted and is a goodsource of water; and KZeinia Zongifzbra is used in Bakgatlarainmaking ceremonies (Hargreaves 1990). An increasingdemand for some of these species could have detrimentaleffects on the wild populations. As there are no lawsprotecting the flora of Botswana (see Existingconservation measures below) some plants may also beremoved from the wild by avid collectors. A number ofspecies are also affected by other human activities, such asexpanding urbanisation, mining, and agriculturalactivities, particularly the planting of crops and grazing.

NamibiaDue to the low human population density in most ofNamibia and the relatively large area covered by naturereserves (13 per cent according to Maggs et al. 1994), onecan argue that there is little threat to the widespreadsucculent species. The situation is, however, somewhatdifferent for those succulent species of more restricteddistribution, particularly the local endemics. These speciesare largely found in the Namibian centres of plantendemism (see Maggs et al. 1994) where overexploitationof land can easily lead to the endangerment of locally orregionally endemic species. In Damaraland and theKaokoveld very intensive exploitation of certain plant

In the coastal mining area of Diamond Area No. 1(the coastal strip between Luderitz and Oranjemund)large scale destruction of the area immediately adjacentto the coast has taken place over the last sixty years. Fewbotanists have been given permission to enter the miningarea proper, and hence one can only speculate about theextent of the damage and the effect of this on thevegetation and the loss of species diversity, particularlyamongst the succulent plant species. From the work doneby Jurgens on the vegetation dynamics of the sandveldarea to the south of the Orange River, one can predictthat a similar pattern would be found further to the north,expecting under natural conditions a situation with highspecies turnover and rapid dynamics. The mostly sandysoils, comprising largely wind blown material, haveresulted in the evolution of species with the ability toregenerate fairly quickly. However, the large scale andrapid nature of the destruction caused by the miningoperations will have certainly resulted in the extinction (atleast locally) of some species. The mining activities andrehabilitation plans for the area do not incorporateappropriate botanical expertise; this is largely becauseonly Consolidated Diamond Mining personnel areallowed to work in the mining area itself, very few ofwhom have the appropriate training. Perhaps of greaterconcern than the mining activities, which are largelyconfined to the coastal strip, are the recreational activitiesof the local inhabitants, particularly four-wheel drive off-road driving (Maggs et al. 1994).

Along the lower Orange River valley (betweenHunsberge/Richtersveld and the Atlantic Ocean),diamond mining activities have increased markedly duringthe last ten years. Large stretches of this extremely aridlandscape have been destroyed by surface mining and thedeposition of screened material (mine dumps). Jurgens(pers. comm.), who has a number of permanent plots inthis area, has found that extremely long time scales arerequired for the regeneration of the destroyed vegetation.

72

This area, termed the West Gariep Circle, is a local centreof endemism (Jurgens 1991) which has been very poorlyresearched as a result of restricted access to the area(Sperrgebiet). Mining occurs on both sides of the OrangeRiver, thus resulting in the destruction of vegetation andloss of species in both Namibia and South Africa. Hereagain, the exploitation techniques used do not take intoconsideration the dynamics involved in the regenerationof vegetation.

Succulent collectors also cause some destruction inNamibia, particularly in the Hunsberge area, whereextremely rare species like TyEecodon singularis o rattractive and highly sought after species likePachypodium are ‘hunted’. Of great concern are recentreports of wild-collected plants removed from therestricted diamond mining area of the Sperrgebiet beingoffered for sale in Europe (D. Newton, pers. comm.;Maggs et al. 1994). Progressive opening up of the moreremote parts of southern and western Namibia formining, farming, and tourism will undoubtedly result inincreasing losses of plants in the populations of rare andendangered succulent plant species.

LesothoTalukdar (1994) presents a review of the botanicaldiversity of Lesotho and its conservation. Of the threats tothe flora which Talukdar lists, the ones most likely to havean impact on succulent species include: a) expandinghuman settlements; b) an increasing dearth of arable landin the lowlands resulting in intensive cultivation of thefoothills and some mountain areas, often on steep slopesunsuitable for agriculture; c) overstocking - it isestimated that there are currently 2.5 to 4 times thenumber of domestic animals than the grassland is capableof sustaining in good health; d) overgrazing resulting fromoverstocking causing degradation of pasture, loss oftopsoil through removal of vegetation cover and soilerosion, and the loss of species; and e) inundation ofmore than one-third of Lesotho’s total area once theLesotho Highlands Water Project is completed. Thisproject, together with its related activities of roadconstruction etc., will certainly have an impact on thepopulations of Aloe polyphylla and Delosperma ashtonii. Inthe case of A. polyphylla, the greatest threat is theuprooting and sale of this popular spiral aloe. Despite theprotected status of this species (see Legislation below) theremoval of plants from the wild for sale to collectorscontinues unabated (Donnay and Meyer 1981; Talukdar1983, 1994). This is possibly a case where too muchpublicity has been to the disadvantage of the species, as ithas only made the plant more desirable (Talukdar 1994).

South AfricaMuch has been written about the threats to the botanicaldiversity of South Africa in general (see chapters inHuntley 1989 and 1994), and in particular on some of thecausal factors responsible for the rate of decline in the

numbers of species (see Everard 1988; Fourie 1084; Hall1987; Hall et al. 1984; and Nolte 1992). Although naturalcauses, such as climatic shifts, and genetic declinecertainly account for some of the changes in speciesabundances and distribution, it is undoubtedly themodification and transformation of the landscape bypeople which is having the greatest impact, not only onlocalised succulent species but also on widespread species.In summary, these transformations include:

1)

2)

3)

Industrial development, urbanisation, expanding ruralsettlements and their allied transport infrastructure.These activities have often taken place in areas of highdiversity and endemism, particularly along the coast,affecting considerable numbers of rare species.The invasion of natural areas by introduced alienplants which often out-compete and replace theindigenous flora (see chapters in Macdonald ct al.1986).Approximately 70 per cent of southern Africa isutilised by domestic livestock for grazing purposes(Macdonald 1989). Much of the land has been, and inmany places still is, overstocked far beyond its carryingcapacity resulting in overgrazing. This overgrazingtogether with trampling results in the loss ofvegetation cover, increased soil erosion and ultimatelythe loss of species. In addition some succulent species

Vendors selling their wares in the medicinal plantmarket of Ezimbuzini near Du rban, South Africa.

73

4)

5)

6)

7)

8)

are actively eradicated from farms, e.g. many of theTylecodon species are dug out, because they causestock illness (krimpsiekte) and even death when eaten.Other species like Euphorbia aggregata, are viewed byfarmers in the Eastern Cape as nuisance plants (Smith1994).Large portions of the country are planted under cropsresulting in monoculture landscapes with a fewremnant patches of natural vegetation on hill tops orareas unsuitable for ploughing such as along rivers andstreams. Much of the Western Cape for example is apatchwork of wheat fields and vineyards, and manyspecies of Haworthia are confined to road verges orsmall remnant patches of natural vegetation in thesouthern parts of this region. The continued survivalof some of these species is entirely dependant on thewhims of the local farmers and landowners.Extensive afforestation with alien timber species suchas pines, blue gums and wattles, is resulting inconsiderable loss of natural habitat for many succulentspecies.Expanding urbanisation and agriculture both requireincreasing quantities of water. The building of dams tomeet this demand has resulted in the flooding ofvalleys with the loss of plant populations and perhapseven the extinction of some species. No information isavailable as to how many succulents have beenaffected by inundation.Mining for minerals and quarrying for buildingmaterials has a major impact as many succulentspecies are confined to these unusual substrates.Unfortunately, the laws governing mineral rights takeprecedence over conservation laws because of theeconomic importance of the minerals to the country,especially in terms of foreign exchange. Only throughintense public pressure and considerable lobbying ofthe government can mining activities be stopped.Removal of succulent plants from the wild is alsohaving an impact on the continued survival of manyspecies. The full extent of this impact has yet to bedocumented although a number of trade surveysconducted in Europe, Japan, and the USA reveal thatwild collected plants are being exported from southernAfrica (see Fuller and Fitzgerald 1987; Jenkins 1993;Oldfield 1991, 1993). The collectors can be classifiedinto three groups: amateur hobbyists, commercialcollectors, and herbalists. Although some amateursundoubtedly collect plants and seeds from the wild,they seldom do this on a regular basis or remove largequantities of material, to be of any major concern.Commercial collectors, on the other hand, have a fargreater impact as they dig out hundreds andsometimes thousands of plants, even removing wholepopulations. The number of prosecutions over theyears and the evidence of wild-collected plants intrade on the overseas markets indicates that this is anongoing problem.

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Adenium multiflorum, a species frequently wild-collected and sold on the roadsides in Zimbabwe.

The international trade in southern African succulentspecies is being investigated by the South/Eastern Africanbranch of TRAFFIC (D. Newton, pers. comm.). Somesucculent species are collected for their edible, medicinal,or magical properties. Haworthia 1imifoZia for example, isremoved from the wild in large quantities by traditionalherbalists, for sale to people in towns and settlements.The only succulent with medicinal properties which isexploited commercially at present is Aloe f&ox (seeOldfield 1993). The leaves of A. feyox are harvested toextract a mucilaginous gel and bitter exudate known as‘bitter aloes’ which is used by the pharmaceutical andcosmetic industries. A report on this harvesting and itsimpact on the species is being prepared by TRAFFIC(Newton 1993).

SwazilandIn Swaziland, as withthe main threat to

most of the rest of southern Africa,the succulent species is habitat

degradation and destruction due to human activities,mainly from expanding settlements and agriculturalactivities. At least one species, namely Haworthia linz(folia,is collected extensively for use as a traditional medicine.

ZimbabweIn Zimbabwe many of the habitats where the speciallyprotected plants (see National Legislation) occur areseriously endangered or at risk. The principal risks ariseout of mining development, the construction of roads anddams, the construction of high voltage power lines, andthe settlement or resettlement of people (often largenumbers) in environmentally sensitive areas containingnumbers of threatened or high risk species (Kimberley1991).

Collectors do not pose a major problem in Zimbabwesince the number of specialist collectors and cultivators inZimbabwe is very small (Kimberley 1991). There is,however, a threat from the activities of itinerant plantsellers who hawk specially protected indigenous plantspecies at all the major shopping centres in Harare quiteopenly and with impunity (Kimberley 1991). Similarly,specimens of Adenium muZtifZorum are often openlyoffered for sale on road sides to passers-by (Kimberley1991). Every specimen sold undoubtedly causes the sellerto remove further specimens from the wild thus reducingthe size and viability of these populations.

A number of Zimbabwe’s succulents are utilised fortraditional medicinal purposes (see Drummond et al.1975). The demand for traditional medicines hasincreased as the population has grown. These demandscombined with the various environmental changesdescribed above will inevitably result in a number ofsucculent species becoming scarce and possibly evenextinct. The same is probably true for other southernAfrican countries, particularly South Africa andSwaziland.


International legislationSeveral international conventions regulate domestic andinternational trade in protected succulent plants. Themost important is the Convention on International Trade

in Endangered Species of Wild Fauna and Flora (CITES)(De Klemm 1990). The Convention, which came intoforce in 1975, has been ratified by over 130 countriesincluding the southern African states of Botswana,Namibia, South Africa, and Zimbabwe. Swaziland is not asignatory and Lesotho, although a signatory, has not asyet ratified the Convention. A number of southernAfrican succulent and semi-succulent species and generaare listed in Appendices I and II of CITES (Table 3.8).None are listed on Appendix III. Trade in more than 498(14.7 per cent) of southern Africa’s succulent taxa isregulated by the Convention. CITES not only regulatesinternational trade in the listed taxa but also in theirrespective parts and derivatives given various exceptionsdepending on the taxon concerned (see Newton andBodasing 1994 for details on the South African planttrade).

The succulent species in trade varies considerablyfrom year-to-year and is dependent on a number offactors such as the availability of plants, taxonomic statusof the group, availability of good literature on the group,rarity, ease of cultivation, and the popularity of the group.As trends in the succulent industry change, a species indemand today may not be so tomorrow. CITES attemptsto remain abreast of the trends by reviewing listing at ten-year intervals. The Convention of the Parties which meetsevery two years, also reviews and votes on proposals to listor downlist taxa. As mentioned above, TRAFFICSouth/Eastern Africa is conducting a survey (1994195) tosee which southern African succulent species arc in trade.Although it is difficult to obtain exact figures on thenumbers of plants traded, the survey informationcombined with data from field studies will hopefullyindicate which species, if any, are threatened by the trade.Initial results show that almost 2600 taxa of southernAfrican succulents are currently being offered for sale innurseries, indicating that the total number of species ininternational trade is significantly higher than reflected by

Table 3.8 Southern African succulent and semi-succulent taxa listed on the CITESAppendicesThe figures in brackets indicate the approximate number of species or taxa indigenous to southern Africa covered by the listing(Note: Pachypodium namaquanum, formerly listed on Appendix I, was downlisted at the recent 1994 Convention of the Partiesto Appendix II in order to facilitate trade in artificially propagated plants.)

Family Appendix I Appendix I I

Aloaceae

ApocynaceaeAsclepiadaceaeCactaceaeEuphorbiaceaePortulacaceaeWelwitschiaceae

Aloe a/M/a (Stapf) ReynoldsA. pillansi L.GuthrieA. polyphylla Schonland ex PillansA. thomcrolfii Pole-EvansA. vossii Reynolds

All Aloe spp. (196 taxa)

All Pachypodium spp. (5 spp.)All Ceropegia spp. (67 taxa)All Cactaceae species (1 sp.)All succulent Euphorbia spp. (199 spp.)All Anacampseros spp. (29 taxa)Welwitschia mirabilis Hook-f.

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the CITES trade statistics (Newton and Bodasing 1994).The results of this survey will be used to makeconservation recommendations and some additionalspecies and/or genera may well be proposed for listingunder CITES at a future meeting of the Parties.

In addition to the fact that two southern Africancountries are not parties to the Convention, all of thesouthern Africa members are contravening the terms ofthe Convention in that they do not have separatemanagement and scientific authorities. In South Africa atpresent, although the Department of Environment Affairs(aided by the Department of Foreign Affairs) is theofficial management authority, responsibility for enactingCITES is devolved to the provinces. South Africatherefore has four management authorities and all four ofthese provincial conservation agencies also act as thescientific authorities. As can be seen from the provinciallegislation described below, there are many discrepanciesbetween the provinces as to how CITES is interpretedand enacted. In addition, the South African regulationsfor the inspection of plant exports need to bestrengthened with regard to expert official viewing ofsamples of the species being exported, as at present this iseither very superficially done or not done at all (Cowlingand Olivier 1992). The division of South Africa into ninenew provinces requires that the issue of managementauthorities and regulations be addressed.

Following the Earth Summit in Rio de Janeiro in June1992, all the countries in southern Africa becamesignatories to the Convention on Biological Diversity. Theobjectives of this Convention include “the conservation ofbiological diversity, the sustainable use of its componentsand the fair and equitable sharing of the benefits arisingout of the utilisation of genetic resources” (Article 1,UNEP 1992). In terms of the Convention, ContractingParties are required to “develop national strategies, plansor programmes for the conservation and sustainable useof biological diversity or adapt for this purpose existingstrategies, plans or programmes” (Article 6, UNEP 1992).Although a number of southern African countries stillhave to ratify the Convention, and those who have doneso still have to start preparing a national strategy, theConvention has significant implications for theconservation of southern African succulent plant species.

The only regional treaty of international relevance isthe 1968 African Convention on the Conservation ofNature and Natural Resources (the African Convention).Under this Convention, the semi-succulent IKeZwitschiamirabilis is the only succulent species to be listed for fullprotection. In addition, Swaziland is the only southernAfrican country which is a party to this Convention.

National (country and provincial) legislationEach country within southern Africa has numerous legalstatutes which provide for the protection of the formallyproclaimed conservation areas such as national parks,game reserves, nature reserves, forest reserves, water

catchment areas, wilderness areas, etc., and for the flora,fauna, and historical objects they contain. In addition tothese statutes, there are a number of laws which protectindividual species of indigenous plants outside of thedeclared protected areas. It is this legislation which is ofmost relevance to the conservation of succulent species.The relevant laws and the succulent species they protectare briefly discussed here on a country-by-country basis.

BotswanaThere are no specific protection laws for plants inBotswana (Hargreaves 1992; Moyo et al. 1993) althoughplants are protected in the national parks, game reserves,sanctuaries, wildlife management areas, controlledhunting areas and forest reserves by the WildlifeConservation and National Parks Act No. 28 of 1992 andthe Forest Act No. 29 of 1980. In terms of the NationalParks Act any person found in or leaving a protected areain possession of an object of “scientific interest”, e.g. asucculent plant, has committed an offence. Unfortunatelyvery few of the conservation areas are in the southern andeastern parts of the country where most of the succulentspecies occur. In terms of the Forest Act a number ofspecies are declared protected trees. However, under theForest (Declaration of Protected Trees) Order of 198 1(Section ll), those species which occur on state land, e.g.Adansonia digitata (a semi-succulent), may be utilised bythe inhabitants of specified villages/towns or habitations.In order to provide some protection for severelyendangered species, the National Museums of Botswanais investigating the possibility of declaring some of thesites where these species occur as “National Monuments”under the Monuments and Relics Act 15 of 1970 (theAloe marlothii forest at Molepole will be the first testcase). In terms of this act, a monument may be defined as“any area of land containing rare or distinctive orbeautiful flora”. However, even if this measure issuccessful it can only provide protection for a fewindividual specimens, not all the plants of a species.Therefore, there is an urgent need for the enactment andenforcement of suitable legislation to protect the flora ofBotswana.

LesothoThreats to the survival of the national flower of Lesotho,Aloe polyphylla, prompted the development of nationallegislation to protect this and other threatened species. In1938 a Proclamation was issued by the ResidentCommissioner of Basutoland, prohibiting the removal,export, sale, or destruction of A. yoZyphyZZa (Talukdar1983). This Proclamation was replaced by the HistoricalMonuments, Relics, Fauna and Flora Act No. 41 of 1967which set out regulations protecting the flora of Lesotho.This Act also made provision for the formation of theProtection and Preservation Commission (PPC) whoseresponsibility it was to enforce the Act. On the advice ofthe PPC the government of Lesotho issued a Legal Notice

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(No. 36 of 1969) in which the Protected Flora of Act 41were defined. The first item in the schedule of protectedflora provides for the protection of all Aloe species, butwith particular reference to A. poZy$yZEa including itsseeds and flowers. Cussonia species were the only othersucculent or semi-succulent group to be listed asprotected flora. The PPC also published a Public Noticein 1970 which prohibited the removal and/or export of A.PolvphyZZ~ (Talukdar 1983). As mentioned above, despiteall these legal attempts to protect this species, protectionhas very rarely been enforced and populations of thespecies have continued to decline (Donnay and Meyer1981; Talukdar 1994). Although this species is unlikely tobecome extinct within the next five years, an integratedconservation plan is required which includes traditionalconservation measures, the involvement of local villagechiefs and headman, combined with an active publiceducation campaign. The flora protection laws, finesimposed, and law-enforcing arm also need to bestrengthened in order to protect other succulent speciesin Lesotho, and the personnel involved must be madeconversant with what is protected and why.

Aloe polyphylla, the national flower of Lesotho and ahighly Endangered species because of wild-collecting.

NamibiaThe Constitution of Namibia makes provision for theconservation of its flora and fauna as part of governmentpolicy by stating that “The State shall actively promoteand maintain the welfare of the people by adopting . . .policies aimed at . . . the maintenance of ecosystems,essential ecological processes and biological diversity inNamibia and utilisation of living natural resources on asustainable basis for the benefit of all Namibians, bothpresent and future...” (Article 95). At present this policy isenforced by the Nature Conservation Ordinance (No. 4 of1975) as amended in 1986 (Act No. 27) and 1990 (Act No.31). Under Schedule 9 of this Act, a large number ofsucculent species are listed as ‘Protected Plants’; these arelisted in Table 3.9. It is interesting to note that the finesand periods of imprisonment for offences relating toanimals have been amended and substantially increased(Act No. 31 of 1990) while those for plants haveremained unchanged. Such disparities between the lawsconcerning flora and fauna are a common feature of allsouthern African countries.

Despite the above legislation there are no adequatemethods of law enforcement in place to protect individualspecies and the list of protected plants is desperately inneed of revision (Maggs et al. 1994). The conservationlaws in Namibia are currently being redrafted, and anenvironmental lawyer was recently appointed as aconsultant to help in preparing the draft (M. Strobach,pers. comm.).

South AfricaWith the establishment of the Union of South Africa in1910, the responsibility for the protection of the flora andfauna was accepted by the four provincial administrations(Section 85, Paragraph 10 of the South Africa Act 1909)and this has remained virtually unchanged since then (seeFuggle and Rabie 1992 and Glavovic 1993). The mostimportant legislation in terms of which indigenous plantsoutside protected areas are conserved are the variousprovincial nature conservation ordinances. The definitionof the term ‘indigenous plant’ as used in the ordinancesvaries, but in general it means any species of plant whichis indigenous to the Republic of South Africa andNamibia, irrespective of whether it is or has beencultivated, or whether it is no longer growing in the wildstate, or has for some time not been growing in the wildstate. The Orange Free State and Transvaal ordinancesfurther specify that protection covers the flower, seed,fruit, cone, bulb, tuber, stem, root or any other part ofsuch plant. The latter also specifies that the plant can bealive or dead. All the ordinances exclude those plantsdeclared to be noxious weeds in terms of any law.

The provincial ordinances will all have to be reviewedand changed as a result of the political changes withinSouth Africa (see discussion below on changes to thelegislation). However, as these ordinances are still inoperation they are discussed here and the succulentspecies they protect are listed in Table 3.10.

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Cape - The Cape Nature and EnvironmentalConservation Ordinance 19 of 1974 provides for theprotection of certain plant species indigenous to theRepublic of South Africa and Namibia (see Table 3.10).Schedule 3 of the Ordinance lists the ‘endangered flora’,which includes those species in danger of extinction andthose listed in Appendix I of CITES, but excludes thosespecies listed in the Appendix which are specified inSchedule 4 (for example, Aloe polyphyh, an endangeredspecies listed on CITES Appendix I, is not included inSchedule 3 because it is included under allAloe species inSchedule 4). Schedule 4 of the Ordinance lists thosespecies declared to be ‘protected flora’ and includes allspecies listed on CITES Appendix II, except for thosespecified in Schedule 3 (Pachypodium namaquanum is anexample of this). Under the Ordinance no person maysell, donate, receive as a donation, pick, or import into,export from, or transport through the Province any‘endangered flora’ without a permit issued by the Directorof Cape Nature Conservation. Similarly no person maypick ‘protected flora’ without a permit or ‘indigenousunprotected flora’ without the written, dated and signedpermission of the land owner or other authorised person.The sale of protected flora is also controlled by theregistration and licensing of flora growers and sellers.

Natal - Under the Natal Nature ConservationOrdinance 15 of 1974, as amended in 1993, Schedules 10,11 and 12 list ‘unprotected indigenous plants’, ‘protectedindigenous plants’, and ‘specially protected indigenousplants’, respectively. No succulent species are included onthe list of ‘unprotected indigenous species’. Schedule 11declares all plants indigenous to the Republic of SouthAfrica and Namibia, except those listed in Schedules 10and 12, to be protected. This list therefore includes thevast majority of succulent taxa listed in Table 3.4, exceptfor those specified in Schedule 12 and 12A. The succulenttaxa listed as specially protected plants on Schedule 12are listed in Table 3.10. Schedule 12A refers to all plantson CITES Appendix I and Appendix II. In terms of theOrdinance, the Natal Parks Board is charged with theconservation of indigenous plants in Natal and it is theirresponsibility to issue permits or licenses which governthe picking, sale, possession, import, or export of‘protected indigenous plants’ and ‘specially protectedindigenous plants’.

Orange Free State - In the Orange Free State, Section30 of the Nature Conservation Ordinance 8 of 1969declares all the species specified in Schedule 6 which areindigenous to the Republic of South Africa and Namibiato be protected plants. The succulent species specified arelisted in Table 3.10. In terms of the Ordinance no personmay pick any ‘protected plant’ without a permit issued bythe Administrator and no person may pick any indigenousplant without the written permission of the land owner. Inaddition, no person may sell, donate, import into or

export from the Province any ‘protected plant’ or a plantof an ‘endangered species’ (CITES Appendix I species) or‘scarce species’ (CITES Appendix II species), exceptunder the authority of a permit from the Administrator.

Transvaal - The Transvaal Nature ConservationOrdinance 12 of 1983 has two Schedules providing for theprotection of plant species indigenous to the Republic ofSouth Africa and Namibia. Schedule 11 (Section 86(l)(a)) specifies ‘protected plants’ and Schedule 12(Section 86 (l)(b)) p fs eci ies ‘specially protected plants’.Schedule 11 excludes all plants which have been improvedby selection or cross-breeding, and it excludes all speciesofAloe which do not occur in the Transvaal and a numberof relatively common Aloe species (see Table 3.10 forexcluded species). The succulent species declared to be‘specially protected plants’ are listed in Table 3.10. Interms of the Ordinance no person may possess, pick, sell,purchase, donate or receive as a donation, import into orconvey within, or export or remove from the Province a‘protected plant’ or ‘specially protected plant’ unless theyhave a permit authorising them to do so. No indigenousplants may be picked within a nature reserve without apermit nor on private land without prior writtenpermission from the land owner or occupier. All nurseryowners are required to be registered in order to sell anyspecies of protected plant cultivated in their nurseries.

In addition to the ‘protected’ and ‘specially protected’plants, all species listed on Appendices I and II of CITES,and any readily recognisable part or derivative thereof, isdeclared by the Transvaal Ordinance to be an endangeredspecies or a rare species respectively. No person mayimport into or export or remove from the Province anendangered species or a rare species, unless they have apermit authorising them to do so.

Adequacy of provincial legislationThe current legislation for the conservation of SouthAfrican flora is deficient in many respects:

1)

2)

3)

4)

There is no formal national policy on indigenous flora,a situation which is exacerbated by divided control andthe lack of prescribed co-ordination between thevarious conservation agencies.It is clear from the above provincial ordinances thatthe existing laws are fragmented, haphazard, and ofteninconsistent with each other, particularly concerningwhich plant species (even which parts of plants) areprotected and what level of protection they havewithin various parts of South Africa.There are a number of loopholes within the legislationwhich are easily exploited by unscrupulous plantdealers, particularly with regards to the transfer ofplants between provinces.Most of the ordinances are fairly old and as a resultthe taxonomy of the taxa listed in the schedules is out-dated.

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5)

6)

7)

The laws concerning plant species are generally farless stringent than those for animals, particularly forlarge mammals like elephant and rhino (this is true forall southern African countries).Levels of enforcement vary considerably fromprovince to province, and in many cases fall far shortof what is desirable. This is not a reflection of theabilities of the law enforcers themselves but of theirsmall numbers, budgetary shortages, and vast areassubject to their control (Cowling and Olivier 1992).Even if an offender is apprehended and prosecuted,the fines imposed are often too low to act as a seriousdeterrent to commercially-minded plant collectors(Hilton-Taylor and Le Roux 1989).

The election of a new government, the development of anew constitution, the reincorporation of the formerhomeland areas into South Africa, and the division ofSouth Africa into nine new provinces since April 1994 hasmany implications for conservation. All provinciallegislation, including the conservation ordinances, willhave to be either redrafted or drafted from new. Thispresents an ideal opportunity for the conservationcommunity to become involved in the legal processeswhich provide protection for our flora and fauna. Ideally,there should be a national conservation statute withregional administration of it. If it is decided to maintainthe present system, all disparities and inconsistenciesbetween provinces must be removed, when theconservation ordinances are drafted or redrafted. Theschedules of protected plants need to be recompiled andupdated on a regular basis based on good scientificinformation (the national status of a species must beconsidered, not just its local provincial status). The finesfor offences must also be adjusted so that they act as aneffective deterrent. Adequate funding must be madeavailable by government to ensure the efficient enactmentand policing of this new legislation. At the same time,however, the new legislation will have to make allowancesfor aboriginal rights and the harvesting practices ofindigenous people. It will be a long time, if at all, beforethe tribal cultures are assimilated into western society,hence their needs and traditional values have to beaccommodated within the legal system. Controlled accessto natural resources (including succulent plants)consistent with traditional harvesting practices and theprinciple of sustainable utilisation will have to be allowed.Although better control of trade in plants used fortraditional healing and other traditional uses will beessential, the law must be sensitive to the needs of thepeople involved in such trade.

SwazilandThere is an assortment of legislation in Swaziland relatingto the protection of its indigenous flora. The NationalTrust Commission Act of 1972, which created theSwaziland National Trust Commission to oversee nature

conservation and the preservation of Swazi heritage,provides legislation concerning protected areasproclaimed under this Act (Braun and Prendergast 1992).For succulent species which occur outside of theprotected area network, the Flora Protection Act No. 45of 1952 (as amended by Government Notices 146 of 1974and 32 of 1975) is of most importance. Under this Act anumber of species are listed in a Schedule as ‘protectedindigenous plants’. The succulent species on the Scheduleare listed in Table 3.9. In terms of the Act ‘protectedindigenous plants’ means any plant, shrub or tree (allparts included) indigenous to southern Africa which isincluded in the Schedule. The Act prohibits any personfrom plucking, gathering, cutting, uprooting, injuring,breaking, destroying, selling, exporting, or possessing any‘protected indigenous plants’ unless they have a permitfrom the Minister, or the plants have been specificallycultivated, or the land is required for cultivation orconstruction. In terms of the Act the Minister ofAgriculture is enabled to establish one or moreindigenous floral reserves.

Attempts were made in 1975 to amend the FloraProtection Act by means of a draft Amendment Order.The aim of this amendment was to make provision forbetter and more effective protection of the flora ofSwaziland, and to increase the penalty for anycontravention of the Act. Many of the clauses in the draftwere unfortunately not accepted. Enforcement of theconservation legislation in Swaziland, as in most ofsouthern Africa, is extremely difficult and many offencesgo unprosecuted (Braun and Dlamini 1994).

ZimbabweA number of plant species are specially protectedindigenous plants in terms of the Zimbabwean Parks andWildlife Act of 1975. The succulent species which areafforded protection by this Act are listed in Table 3.9.Under this Act, it is a criminal offencc to dig up, pick,tamper with, or remove ‘specially protected indigenousplants’ from habitat.

Land owners were exempt from the provision of theAct until 1985 when an amendment to the Parks andWildlife Act of 1975 inserted a new Section 46A into theAct which empowers the Minister of the Environmentand Tourism, if considered necessary or desirable to do soin the interests of preservation, conservation, propagationor control of any indigenous plants within Zimbabwe orany area of Zimbabwe, to prohibit (absolutely or subjectto conditions, indefinitely or for a specified period) theland owner from picking or selling specified indigenousplants.

It is also an offence to sell a ‘specially protectedindigenous plant’ except in terms of a permit issued by theMinister. Likewise, it is an offence to purchase a speciallyprotected indigenous plant except from a permit holderor an authorised dealer in such plants.

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Table 3.9 Succulent plants declared to be protected species in Namibia, Swaziland, andZimbabweIn some cases, the names of the taxa listed in the Acts have gone into synonymy; the currently accepted names are shown herewith the synonyms in brackets or, if the species of a genus have been transferred to a number of genera, these genera are givenin brackets, Genera followed by 3pp.’ include all species.

Namibia

AizoaceaeAridaria noctifloraAstridia spp.Cephalophyllum spp.Chasma tophyllum musculinumCheiridopsis spp.Conophytum spp,Dinteranthus spp.Ebracteola spp.Fenestraria rhopalophylla ssp. aurantiacaHereroa spp.Jensenobotrya lossowianaJuttadinteria spp.Lapidaria margaretaeLithops spp.Nananthus aloidesOpthalmophyllum spp. (many of these are now

Conophytum spp.)Psammophora spp.Ruschia spp.Schwantesia spp.Stoeberia spp.Titanopsis spp.Trichodiadema spp.AloaceaeAloe spp.Chortolirion angolense (= C. bergerianum)Gasteria pillansii var. ernesti-ruschiiHaworthia venosa ssp. tesselata (=H. tesselata var. engleri.)

ApocynaceaePachypodium lea/iiP. namaquanumAsclepiadaceaeCaralluma spp. (transferred to either Quaqua or

Pachycymbium)Ceropegia spp.Duvalia spp.Hoodia spp.Huernia spp.Huerniopsis spp.Piaranthus spp.Stapelia spp.Tavaresia barklyi (= Decabelone barklyi)Trichocaulon spp. (transferred to either Hoodia or Lavrania)CrassulaceaeAdromischus spp.Crassula spp.MoringaceaeMoringa ovalifoliaPortulaceaeAnacampseros spp,Portulacaria pygmaeaVitaceaestem succulent Cyphostemma spp.WelwitschiaceaeWelwitschia mirabilis

Swaziland

Aizoaceae Pachypodium saundersiiLithops spp. Brachystelma spp.Frithia spp. Caralluma spp. (transferred to Pachycymbium)Aloaceae Ceropegia spp.Aloe spp. Duvalia spp.Haworthia spp. Huernia spp.Apocynaceae Stapelia spp.Adenium multiflorum Stultitia spp. (transferred to Orbea, Orbeopsis orA. swazicum Pachycymbium)

Zimbabwe

AloaceaeAll species and varieties of Aloe, including the natural hybridsApocynaceaeAdenium multiflorum (=A. obesum) ssp. multiflorumPachypodium saundersiiHoodia lugardiiTavaresia barklyi

EuphorbiaceaeEuphorbia davyiE. deciduaE. memoralisE. wildiiPassifloraceaeAdina fructicosaA. spinosa

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Protected area networkA large number and variety of protected areas such asnational parks, game reserves, state forests, wildernessareas, local authority nature reserves, bird sanctuaries,water catchment areas, etc., have been established withinthe countries of southern Africa. The aim of most of theseareas is principally to ensure that entire ecosystems,communities, or habitats are conserved, rather than justindividual plant or animal species. A few reserves havebeen proclaimed specifically for the preservation of asingle species (mainly mammals as for example in the caseof the Addo Elephant National Park or BontebokNational Park), but in so doing much of the naturalhabitat is also conserved. The protected area network ineach southern African country is described and reviewedby WCMC (1991) and additional details concerning thepreservation of key South African ecosystems are given inHuntley (1989). The adequacy of the existing protectedarea network in relation to the preservation of the flora ofeach southern African country is reviewed in Huntley(1994). From that information it appears that some 30million hectares (9.8 percent of the area of southernAfrica) is included in state or semi-state conservationareas. This figure would be considerably higher if onewere to include all the privately owned and local authorityreserves, but as they have no long-term conservation value(in terms of current legislation) they have been ignored.

Many of the protected areas in southern Africa havebeen declared on an ad hoc basis or have been the resultof the focus on suitable extensive areas for theconservation of large mammals, rather than the result of apreconceived design for maximising the preservation ofbiological diversity (Huntley 1978; Siegfried 1989). Giventhat six percent of South Africa is set aside forconservation, it is surprising to find that possibly 74

percent of South Africa’s vascular plant species occurwithin the reserve system (Siegfried 1989). This resultmay be an artefact of the data used for the analysis,although the finding is supported by data for Swazilandwhich indicates that 60 percent of Swaziland’s plantspecies are protected within two small nature reservescovering only two percent of the total area of that country(Braun and Dlamini 1994). Siegfried (1989) was only ableto obtain complete plant checklists for 52 of SouthAfrica’s reserves and partial checklists for 153 reserves,whilst 377 had no lists at all. This situation is still true formuch of southern Africa today and as a result it isimpossible to say at this stage how many succulent speciesoccur in the protected area network. However, given thefacts that succulents are often highly localised toparticular and often unusual habitats and that the areaswith the highest diversity of succulents are the leastconserved (see Hilton-Taylor and Le Roux 1989; Cowlingand Hilton-Taylor 1994) it is reasonable to conclude thatvery few of our succulent taxa are conserved in theprotected area network. The Succulent Karoo forexample, an area with the greatest diversity of succulentplants in southern Africa, has less than two percent of itsarea set aside for conservation (see Hilton-Taylor and LeRoux 1989; Hilton-Taylor 1994).

In an attempt to identify the gaps in the southernAfrican (excluding Zimbabwe) protected area network,Rebel0 (1994) used iterative selection procedures toanalyse data from PRECIS (the PRE (NationalHerbarium, Pretoria) Computerised Information System).The main aim of this analysis was to determine theoptimal placement of nature reserves in relation to thecentres of endemism. Rebel0 (1994) has shown that 93percent of southern Africa’s plant taxa can be conservedoptimally in just ten percent of southern Africa’s area,

The Mathews Rockery atKirstenbosch NationalBotanical Garden whichcontains a large varietyof Aloe species andmany species ofAizoaceae.

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The new arid plants conservatory at KirstenboschNational Botanical Garden will display to the publicone of the largest collections of southern Africansucculents in the world.

require 22 per cent of the area. Comparing the existingprotected area network to the optimal reserveconfiguration, Rebel0 (1994) has indicated in which areasland needs to be set aside for conservation. The largestcontiguous area requiring preservation that currently hasthe fewest reserves is the Succulent Karoo as discussedabove. There are a number of problems with the PRECISdata set, including incompleteness (especially with regardsto succulent taxa), incorrect data, uneven coverage(especially for the arid regions), and the coarse scale usedfor recording distributions (Rebel0 1994). In order toassess how effective the current reserve network is inpreserving succulent plants and to identify gaps in thenetwork, detailed distribution information would have tobe obtained from all the main herbaria in the region andsubjected to a similar analysis.

Although approximately 14 per cent of Zimbabwe hasbeen set aside for conservation purposes, the completerange of vegetation types is not adequately conserved andthere are many rare and/or endemic plant species whichare not protected in the existing network (Timberlake andMuller 1994). It is also alarming to note that 11 of the 25Botanical Reserves, established in 1975 under the Parksand Wildlife Act, were deproclaimed in 1979 presumablybecause of partial destruction or degradation (Timberlakeand Muller 1994). Using a hierarchical approach,Timberlake and Muller (1994) have identified a numberof areas important for the conservation of botanicaldiversity in Zimbabwe. They also stress that simplyidentifying these areas is not sufficient for conservation.Land-use planning and appropriate managementtechniques need to be developed as well (Timberlake andMuller 1994).

Ex situ conservationAlthough regarded as the ideal option, there are

significant drawbacks to ivt situ conservation of plantspecies in protected areas. These areas are vulnerable to

natural disasters such as pests, fires, and storms;vulnerable to political whims and ever-increasingpressures for land; rapidly becoming more expensive tomanage and protect; protect only a limited number ofspecies and genotypes; and susceptible to the effects ofglobal warming (Moss 1994). A complementary tool to insitu conservation is the use of ex situ gene bankingmethods. Gene banking includes the preservation offrozen seeds in conventional seed banks, the growing of

5 plants in field gene banks such as in botanical gardens,$ arboreta and nurseries, and in vitro techniques such as$ tissue culture for micropropagation and cryopreservationF (Hawkes 1992). The conservation of species by means ofcj ex situ techniques also has many disadvantages, for

example: only a very small portion of the total gene poolcan be stored; it is a very expensive and labour intensiveoperation; evolution is effectively stopped; genetic shiftand drift may occur in multiplication cycles, thuscompromising genetic integrity; and very little is knownabout the long-term viability of seed of most wild species(Moss 1994). On the positive side, however, cx sitrrsamples serve as an insurance policy or hedge againstpopulation extinction or catastrophic loss of habitat andthey can also be used to enhance the probability ofsuccess of the irreplaceable in situ efforts.

EX situ conservation of plant diversity by means ofseed banks has been slowly getting under way in southernAfrica since 1987 with the development of a regionalgenetic resources programme known as the SADC(Southern African Development Community) RegionalGene Bank (see Moss 1994). The main focus of thisprogramme, which involves the establishment of nationalgene banks, has been on genetic resources which haveeconomic value as agricultural crops, forages, medicinalplants, horticultural crops, traditional wild foods, etc.Unfortunately, very few succulent species have beenconsidered to have enormous economic potential towarrant the collection and storage of their seed. Asidefrom a handful of succulent seeds collected during recentexpeditions to Namibia and South Africa there is verylittle seed of southern African succulent species (theremay be some seed of a few semi-succulents) stored in thenational gene banks established so far, and it is doubtfulwhether there are any samples in overseas gene bankssuch as Wakehurst Place at the Royal Botanic Gardens,Kew (see Braun and Prendergast 1992; Prendergast1994). Even the small gene bank established by the BolusHerbarium of the University of Cape Town forendangered species contains no seed of any succulentspecies (A.V. Hall, pers. comm.).

Most of the countries in southern Africa (exceptingSwaziland) have at least one botanical garden. Some ofthese gardens are very new such as the one beingestablished near Gaborone by the National Museums ofBotswana, and that being developed around the NationalHerbarium in Windhoek by the National BotanicalResearch Institute of Namibia. In Lesotho, the University

82

Table 3.10 Succulent plants declared to be Protected Species (*), Specially ProtectedSpecies (#) or Endangered Species (+) by the nature conservation ordinances of the fourprovinces of South AfricaIn some cases, the names of the taxa listed in the Ordinances have gone into synonymy; the currently accepted names areshown here with the synonyms in brackets or, if the species of a genus have been transferred to a number of genera, thesegenera are given in brackets.

CapeAizoaceaeAll species in the family formerly called

the Mesembryanthemaceae”AloaceaeAloe spp.* (except for A. Xerox)Aloe buhrii +A. erinacea +A. pillansii +Gasteria nitida var. nitida * (=G. beckeri)Haworthia spp. *ApocynaceaePachypodium spp I *P. namaquanum +AsclepiadaceaeAll species*

CrassulaceaeCrassula coccinea * (=Rochea coccinea)C. columnaris * (presumably including

the subspp.)C. petfoliata var. minor * (= C. falcata)C. perfoliata var. perfoliata *C. pyramidalis *Kalanchoe thyrsiflora *DioscoreaceaeDioscorea eiephan tipes *

(= Tes tudinaria elephantipes)D. sylvatica var. syivatica * (=T. syivatica)EuphorbiaceaeEuphorbia bupieurifoiia *E. fascicuiata *

E. globosa *E. horrida * (presumably including the 3

varieties)E. groenewaldii #E. meloformis *E. obesa*E. schoenlandii *E. symmetrica *E. valida *PortulacaceaeAnacampseros spp. *All familiesAll species on CITES Appendix I 1All species on CITES Appendix II*

NatalAloaceaeAloe spp.*Gasteria spp. #Haworfhia spp.#ApocynaceaeAdenium spp. #Pachypodium saundersii #

AsclepiadaceaeBrachysteima spp. #Caraiiuma spp. # (Natal species

transferred to Pachycymbium)Duvalia spp.#Huernia spp. #Stapeiia spp.#Stuititia spp.# (transferred to Orbea,

Orbeopsis or Pachycymbium)

DioscoreaceaeDioscorea spp.#All familiesSpecies indigenous to the Republic of

South Africa or Namibia*, species onCITES Appendix I and CITESAppendix II*

Orange Free StateAizoaceaeLithops spp. *Neohenricia sibbetii *Pieiospiios spp. *Titanopsis calcarea *AloaceaeAloe spp.#Haworthia venosa ssp. tesselata * (=H.

venosa ssp. recurva)

Transvaal

ApocynaceaePachypodium spp. *AraliceaeaCussonia spp.#AsclepiadaceaeStapelieae spp. *Ceropegia spp. *Trichocauion spp. * (transferred to Hoodia

or Lavrania)

BegoniaceaeBegonia sutheriandii *DioscoreaceaeDioscorea spp. *EuphorbiaceaeEuphorbia spp. *PortulacaceaeAnacampseros spp. *

AizoaceaeLithops lesiiei # (presumably including the

7 varieties)Frithia puichra # (presumably including

the variety)AloaceaeA/i Aloe * species excluding (a) those not

occurring in the Transvaal and (b) thefollowing: A. acuieata, A. ammophiia,A. arborescens, A. ctyptopoda (=A.wickensii), A. grea theadii var. davyana(=A. barbetfoniae), A. castanea,A. davyana, A. mutans, A, globuligemma,A. grandiden ta ta, A. iutescens,A. mariothii , A. parvibracteata, andA. transvaaiensis

Gasteria spp. *Haworthia spp. *

ApocynaceaeAdenium multifiorum # (=A. obesum)A. oleifolium #A, swazicum #Pachypodium saundersii#AraliceaeaCussonia spp. *AsclepiadaceaeBrachysteima spp.#Ceropegia spp. #Duvaiia spp. #Hoodia currorii ssp. iugardii # (=H.

lugardii)Huernia spp. #Huerniopsis spp.#Orbea spp. #Orbeanthus spp.#Orbeopsis spp. #

Pachycymbium spp.#Stapeiia spp. #BegoniaceaeBegonia spp. #BombacaceaeAdansonia digita ta #DioscoreaceaeDioscorea spp.#EuphorbiaceaeEuphorbia barnardii #E. ciivicola #E. grandiaiata #E. knobelii #E. perangusta #E. restricta #E. rowiandii #E. tortirama #E. waterbergensis #

83

of Roma has a garden in Maseru, and there are at leastfour gardens in Zimbabwe. South Africa has the greatestnumber and diversity of gardens ranging from eightNational Botanical Gardens - under the auspices of theNational Botanical Institute (NBI) whose primary focus isthe growing of indigenous flora - to a large number ofuniversity, municipal, and privately owned gardens.

Practically all of the gardens in southern Africa growsucculent species; however, not all of these gardensmaintain accurate records about the sources of their plantmaterial, the location of this material in their garden, andthe history of these plants, especially the genetic integrityof their progeny. More importantly, few of these gardensmaintain succulent collections with the goal ofconservation of species; most of the gardens are primarilyintended for recreational purposes. All of these gardenscan, however, play an important role in succulent plantconservation (see Heywood 1992). One of the statedobjectives of the National Botanical Gardens (NBI) inSouth Africa is to establish collections of threatenedplants and to promote the ex situ conservation of rare andendangered species. All of the NBI gardens growsucculent plants; however, it is the Kirstenbosch, Pretoria,and Karoo National Botanical Gardens which have thelargest collections, particularly the latter (see EX situconservation in Chapter 2). No attempt has been made sofar to determine accurately how many species ofsucculents in total are in cultivation at these gardens, orhow many plants there are (especially of different geneticorigin) of each species. Some attempts have been made todetermine the number of threatened species being grownin each garden, but there has been no comparisonbetween gardens. It is therefore difficult to determineexactly how many threatened species in total there are incultivation. The new Red Data List of Southern African

Plants (Hilton-Taylor 1996b) provides the opportunity toassess the numbers of threatened succulent species incultivation.

Table 3.11 provides a preliminary assessment of thesouthern African (FSA region only) Aloaceae taxa incultivation at three of the National Botanical Gardens ofSouth Africa (Karoo, Kirstenbosch, Pretoria) in 1994. Theoverall results are very encouraging with 88 per cent and95 per cent of the Aloe and Haworthia taxa, respectively,in cultivation. All of the taxa in the remaining generawhich occur in the region, namely Astroloba (5 taxa),Chortolirion (1 taxon), Gasteria (22 taxa), and Poellnitzia(1 taxon), are also in cultivation in at least one of thethree gardens. Looking at the threatened species, 91 percent of the threatened Aloe taxa and 85 per cent of theth rea tened H a w o r t h i a taxa are in cultivation.Unfortunately, what these figures do not indicate are theexact numbers of plants of each species being grown. Inmany cases this is a single individual or a few plants, oftenof the same clone. So while the results are good in termsof species diversity, in terms of conservation of geneticdiversity the results are very poor. More attention needsto be paid to this problem by the gardens if they wish tocontinue their role in ex situ conservation.

Despite the various drawbacks and problems inmaintaining species in gardens, this method ofconservation is of considerable value as is clearlyillustrated by Foster (1993) in describing how thecollection of Aloaceae is utilised at the Royal BotanicGardens, Kew. The collections of Aloaceae in thebotanical gardens of southern Africa are an importantinternational scientific and conservation resource. ThePretoria National Botanical Garden, for example, has notonly one of the most comprehensive collections ofsouthern African aloes, but includes 139 AZoe taxa from

Table 3.11 The numbers of southern African Aloe and Haworthia taxa in cultivation at theKirstenbosch, Karoo, and Pretoria National Botanical Gardens, South Africa (figures forHaworfhia were not available for Pretoria) in 1994The figures for each garden are indicated separately and then a combined total for each genus is shown. Column one indicatesthe number of plants in cultivation column two the per cent of southern plants? in cultivation, column three the number ofthreatened taxa in cultivation, and column four the per cent of the total number of threatened taxa in that genusb which are incultivation.

In cultiv. Per cent

AloeKirstenbosch NBG 115 7-lKaroo NBG 7-t 44Pretoria NBG 125 77

Total 143 (163a) 88

HaworthiaKirsten bosch N BG 58 50Karoo NBG 101 88

Total 109 (115aa) 95

a Total number of taxa in the FSA region.b Total number of taxa in the region considered to be threatened.

Threatened Per cent

35 6720 3944 85

49 (549 91

15 3833 83

34 (40b) 85

84

east Africa, north-east Africa, and Madagascar, plus fivespecies of Lomatophyllum from Madagascar and theMascarene Islands.

Proposed conservation strategyDespite all the international and national legal

protection measures, despite the existing protected areanetwork, and despite attempts to establish ex situconservation measures, the current development patternsand continued exploitation of resources in southernAfrica are resulting in the increasing depletion of theregion’s succulent plant diversity. To prevent the loss ofthis diversity a multi-faceted conservation strategyinvolving all the stake-holders is required (see Nolte 1992who presents such a strategy for the conservation ofsucculent plants). Incorporated into the Action Proposalsin Chapter 4 are a number of suggestions and actionswhich are required as part of this strategy. The list is by nomeans comprehensive and the actions are not listed in anyorder of priority. Although suggestions are given as towho the key players for each action are, the proposals arenot intended to be prescriptive or binding in any way.Also, as many of the key players have not been fullyidentified or consulted about their involvement, noattempt has been made to cost or timetable any of theproposals. Many of the actions are also not once-offevents, but rather on-going exercises.

The dramatic political transition that has occurred inSouth Africa since the start of the reform programme,initiated in 1990 and culminating in the inauguration ofMr Nelson Mandela as the President of a democraticSouth Africa on 27 April 1994, is having a profoundinfluence on conservation policy within South Africa andwhich may have repercussions for the adjoining states(Huntley 1996). The new government of national unityhas placed the highest priority on its Reconstruction andDevelopment Programme (RDP) to redress the legacy ofapartheid. The RDP, which focuses primarily on socio-economic disparities, will also be addressingenvironmental issues, and policy outlines indicate thatnew approaches need to be developed by the existingconservation agencies (Huntley 1996). The conservationstrategy included in Chapter 4 (certainly for South Africa)needs to be carried out within the constraints of the RDP.An important policy change required by the RDP is theparticipation of local communities in the decision-makingprocess concerning environmental issues such as landtenure and access to natural resources. Such transparentprocedures concerning resource utilisation (Huntley1996) and the development of good neighbour relations inthe management of protected areas (Fourie 1994) willhopefully dispel the view held by most of southernAfrica’s black majority that saving succulent plants (alongwith rhinos, elephants, and other internationally popularspecies) is an indulgent pursuit of the affluent minority.

The author would like to acknowledge B. Hargreaves, N. Jurgens,and M. J. Kimberley for their contributions and review of thissection. I wish to thank the Directors of the former four SouthAfrican Provincial conservation agencies for access to thethreatened plants data held by their organisations. I am grateful toKerry Walter (formerly) and Harriet Gillett (presently) of theThreatened Plants Unit of the World Conservation MonitoringCentre, for providing me with data for other southern Africancountries. I am also extremely indebted to the following peoplewho provided information on specific plant groups (indicated inbrackets), on a variety of different taxa, or for specific regions: F.Albers (Asclepiadaceae); T. Anderson; M.B. Bayer (Aloaceae,Asclepiadaceae); P.V. Bruyns (Aloaceae, Asclepiadaceae,Asteraceae, Crassulaceae, Euphorbiaceae); W. Boyd; K. Braun;P. Craven; J. du Preez; M. Gerbaulet (Portulacaceae); H.F. Glen(Aloaceae); B. Groen (Aizoaceae, Aloaceae); A.V. Hall; S.A.Hammer (Aizoaceae); D.S. Hardy (Aloaceae); H.E.K. Hartmann(Aizoaceae) ; H. D. I hlenfeld (Aizoaceae) ; N. Jurgens (Aizoaceae) ;H. Kolberg; D. Koutnik (Euphorbiaceae); L.C. Leach(Asclepiadaceae, Euphorbiaceae); G. Maggs; U. Meve(Asclepiadaceae); S. Liede (Aizoaceae, Asclepiadaceae); D.J.McDonald (Aloaceae); A. Le Roux; I. Oliver; P. Phillipson; F.Powrie (Geraniaceae); R. Scott-Shaw; G.F. Smith (Aloaceae); R.Stanvliet; M. Struck (Aizoaceae); J. Venter (Aloaceae); D. Viljoen;E.J. van Jaarsveld (Aizoaceae, Aloaceae, Crassulaceae); J. Vlok;P. Vorster (Geraniaceae); G. Williamson; and N. Zimmermann(Aizoaceae). I apologise to anyone who has inadvertently beenomitted from the above list. We are also very grateful to thefollowing people for providing us with the information onAloaceae taxa in cultivation: Hermi Marent and Ernst vanJaarsveld (Kirstenbosch National Botanical Garden); Ian Oliver,Rina Smit and Deon Viljoen (Karoo National Botanical Garden);and Priscilla Swartz (Pretoria National Botanical Garden). Manythanks to Gideon Smith for his comments on the paper and forhis assistance in preparing Figure 3.1.

North AmericaEdward F. Anderson

The continent of North America has a land mass of24,700,OOO km2 (9,500,OOO mi”), which makes up 16.2percent of the earth’s surface. It extends from 14”30’N atthe Mexico-Guatemala border to 70”N latitude near theArctic Circle. Virtually all types of biomes are present,from arctic tundra to tropical rain forest. Topography isalso highly variable, ranging from sea level to nearly 6000m elevation. The countries of Canada, the USA, and TheRepublic of Mexico form the political and geographicboundaries of this continent.

Table 3.12 lists the plant families of North Americawhich contain native succulent members. Some of thesefamilies, such as the Burseraceae and Fouquieriaceae,have relatively restricted habitats, whereas others, such asthe Crassulaceae, occur in virtually all regions of NorthAmerica. Nonetheless, the succulent flora of NorthAmerica tends to be concentrated in the arid regions ofMexico and the south-western USA, and it is within theseareas that the main conservation problems exist.

The Agavaceae, Cactaceae, Crassulaceae, and

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Table 3.12 The main plant families ofNorth America which contain nativesucculent members

No. genera No. species

Agavaceae (sensu lato) 12 360Aizoaceae 3 15Burseraceae 1 12Cactaceae 79 1170Commelinaceae 12 100Crassulaceae 6 >300Dioscoriaceae 1 63Fouquieriaceae 1 11Portulacaceae 5 many spp.

Fouquieriaceae have representatives that are either listedin Appendix I of CITES, the SEMARNAP Diario Oficial(Mexico), or in the US Endangered Species Act. The twofamilies containing the most vulnerable taxa are theAgavaceae and Cactaceae.

North America can be most conveniently divided intotwo regions: 1) Canada and the United States ofAmerica, and 2) The Republic of Mexico. Canadacurrently has no succulent plant populations under threat.Because the USA and Mexico differ so greatly invegetation types, development of conservation programs,and protected regions, the two political units will beconsidered separately.

Succulent plant flora in the United States

Most succulents are located in the western USA, althoughmembers of the Cactaceae are found virtually throughoutthe country, as well as in much of Canada. The genusSedum of the Crassulaceae is widespread in numerousvegetation types in both countries.

The main succulent plant families in the USA are theAgavaceae, Cactaceae, and Crassulaceae. The Aizoaceae,Burseraceae, Commelinaceae, Fouquieriaceae, andPortulacaceae are also present, but not in large numbersas compared to in Mexico. The Agavaceae has five generaand 70 species in the USA. The Cactaceae has 29 genera(2 endemic) and 246 species. The Crassulaceae has 3genera with more than 100 species.

The main vegetation types with succulent plants arethe Sonoran Desert, Mohave Desert, Great Basin(Sagebrush) Desert, Chihuahua Desert , DesertGrassland, Coastal Desert Scrub, and Chaparral. I

Threats

The most serious threats to succulent populations in theUSA are collectors, off-road vehicles, urban development,agricultural development, the effects of mining and roadconstruction, and the removal of forests or other naturalvegetation. Those regions most seriously threatened aredesert areas near major urban centres in California,Arizona, and New Mexico. One such example is theMohave Desert of California in which off-road vehicleshave destroyed extensive areas of native vegetation. The

Sonoran Desert

development of natural gas lines in New Mexico has alsoimpacted several populations of Sclerocactrrs andEchinocereus. The construction of dams, such as the GlenCanyon dam in Arizona, has also led to the loss of someimportant desert habitat. Several large local populationsof S. papyracanthus have been destroyed by urban growthin Albuquerque, New Mexico. The rapid growth of desertcommunities, such as Palm Springs, California, has alsodestroyed large areas of the Sonoran Desert in that state.Agricultural development in desert regions of Arizonaand California have also heavily impacted desert plants.Livestock grazing and the accompanying trampling byhooves has resulted in some localised damage tosucculents in the south-west. Collectors have impactedmany small populations of species in Sclerocactus andPediocactus, especially in the states of Arizona and NewMexico.

Local uses and commercialisation

Succulents are most adversely affected in the USA bycollectors and people involved in arid landscaping andhorticulture. Despite laws regulating their collection,specimens of Carnegiea gigantea (saguaro) continue to beillegally removed from habitat to be sold in thecommercial landscaping trade. Likewise, specimens ofFouquieria spZendens (ocotillo) and numerous barrel cacti

Carnegiea salvage operation, Arizona.

86

(Ferocactus spp. and Echinocactus spp.) are subjected tocollecting from the field for horticulture. In some regionsof Texas the spines of Opuntia spp. are burned off andlivestock allowed to browse on the plants. Throughout thesouth-western USA Native Americans and hispanicscollect the fruits of Opuntia spp., Carnegiea gigantea, andother cacti for food. Populations of Lophophora williamsii(peyote) have also been affected by the extensivecollecting of ‘heads’ or tops by Native Americans forreligious purposes or others for personal use or sale.Some other cacti with similar physical characteristics, suchas Astrophytum asterias, have also been mistakenlycollected by those who wished to use peyote for itshallucinogenic effects.

Existing conservation measuresProtected areasSeveral federal and state parks and monuments have beenset aside in the south-western USA to protect the nativeflora and fauna, as well as other natural phenomena.Those protected areas which have numerous succulentsare:

ArizonaGrand Canyon National ParkOrgan Pipe Cactus National MonumentSaguaro National Monument

CaliforniaJoshua Tree National MonumentAnza-Borrego State Park

New MexicoCarlsbad Caverns National Park

TexasBig Bend National ParkBig Bend Ranch State Park

UtahCapitol Reef National Park

Several Native American (Indian) reservations also haverestrictions on collecting plants.

Legal protection for succulentsSucculents are protected at three levels in the USA. Moststates in the south-west have laws controlling thecollection of plants, particularly cacti and succulents. Thestate of Arizona has an Arizona Native Plant Law whichspecifically protects all Cactaceae, Crassulaceae, andFouquieriaceae. California has a Desert Native Plant Actwhich requires permits, tags, and seals to collect desertplants on public land. Nevada, New Mexico, and Texashave similar regulations or laws requiring permits forcollecting any plant within the state on private or publicland. At present Utah has no plant protection laws.

The second level of protection is the EndangeredSpecies Act, administered by the US Fish and WildlifeService, in which numerous taxa, including several

Echinocereus triglochidiatus, New Mexico.

succulents (see Table 2.1), are listed as threatened orendangered. Their collection (or destruction) in the wildis strictly controlled by this legislation, especially onpublic land.

The third level of protection is CITES (seeControlling the trade), of which the USA, Canada andMexico are member nations. This treaty controls theinternational trade in many succulents, some of which arenative to the USA.

Ex situ conservationEfforts at ex situ conservation are being made by severalbotanical gardens through the coordination of the Centrefor Plant Conservation (CPC), located in St. Louis,Missouri, which includes provision for the long termstorage of seed of endangered plants. The DesertBotanical Garden in Phoenix, Arizona currently hasprimary responsibility for most succulents from the aridregions of the south-western USA and also propagatesthreatened Mexican species. They are presentlypropagating a total of 16 different threatened andendangered succulent taxa as listed in Table 3.13. Otherbotanic gardens are also propagating rare and endangeredsucculents as part of their regular operations. Theseinclude the Ranch0 Santa Ana Botanic Garden,

87

Claremont, California; the Huntington Botanical Garden,Pasadena, California; and the Botanic Garden of theUniversity of California, Berkeley. As noted in Chapter 2,some commercial nurseries are also making a significantcontribution to ex situ conservation efforts.

Nursery developmentMajor commercial nurseries which specialise in succulentsare found in southern California, Arizona, and Texas.Few grow CITES Appendix I plants because ofcumbersome federal regulations. However, millions ofseed-grown cacti and other succulents are produced therefor the world trade.

Main agencies involved in succulent plantconservationThere are both governmental and non-governmentalagencies involved in conservation activities. The primaryfederal agency is the US Fish and Wildlife Service, whichhas primary responsibility for both the US EndangeredSpecies Act and CITES. The US Department of Justice isalso involved in apprehending and prosecuting those whoviolate federal regulations on international trade insucculents.

The CPC is closely involved in coordinatingconservation activities of more than 20 botanic gardensand other institutions in the USA. The Cactus andSucculent Society of America and its numerous affiliatebranches are also advocates of succulent plantconservation. The national organisation has aConservation Committee. The influence of theInternational Organization for Succulent Plant Study(10s) is strongly felt within the USA, not only throughthe efforts of individual members, but also through itsinternational voice in matters concerning succulent plantconservation. The 10s Code of Conduct has also beenwidely distributed throughout the USA.

Assessment of existing conservationmeasuresIn general, effective legislation exists at both the state andfederal levels to protect and control the trade inrecognised rare and endangered succulents.Unfortunately, prosecution and the ensuing litigation areboth time-consuming and costly. Thus, governmentagencies sometimes are forced to use their limitedresources in dealing with more critical issues, such as thedrug trade and illegal immigration. Education on plantconservation is still needed among many groups ofAmericans. For example, a number of property owners inwest Texas have organised the Trans-Pecos HeritageAssociation to prevent what they perceive to be a federaleffort to ‘lock up’ their private land because a rare plantor animal may occur on it. They actively patrol theirfenced property and arrest trespassers who come ontotheir land for any reason. Unfortunately, most of thesepeople are misinformed of what the government can and

Table 3.13 Threatened and endangeredNorth American succulents in cultivationat the Desert Botanical Garden, Phoenix,Arizona, USA(CPC National Collection, CITES listing (Appendix I or II), orthreatened (T) or endangered (E) under the USEndangered Species Act).

Taxon CPC CITES ESAAgave arizonica * I EA. pa rvi flora * IAriocarpus fissura tus var. lloydii IA. kotschoubeyanus IA. retusus IAztekium ritteri ICoryphantha ramillosa * TC. scheeri var. robustispina * EEchinocactus horizonthalonius

var. nicholii * EEchinocereus chisoensis * TE. ferreirianus * IE. schmollii IE. triglochidia tus var. arizonicus EE. viridiforus var. davisii * EEscobaria minima * IE. robbinsorum *

E. sneedii var. sneedii * IFouquieria columnaris IIF. fascicula ta IF. purpusii ILeuchtenbergia principisMammillaria pectinifera IM. plumosa

4 M. solisioides IObregonia denegrii IPachycereus militaris IPelecyphora aselliformis IP. s tro biliformis ISclerocactus erectocentrus * IS. mariposensis IS. papyracanthus ITumamoca macdougalii * ITurbinicarpus klinkerianus IT. lophophoroides IT. schmiedickeanus IT. schmiedickeanus var. sc hwarzi i I

cannot do on private land. In fact, the avid protection oftheir land from trespassers is also protecting thesucculents that grow there from collectors.

Conservation education must be more creativelypresented to the public and in schools; for too manypeople the terms ‘conservation’ and ‘environment’ arepejorative ideas rather than goals to be sought. Economicimpacts of the Endangered Species Act have tended topolarize people against long-term protective efforts toconserve threatened or endangered species. Often suchfears, though with a real economic basis, are founded on alack of comprehens ive unders tand ing of theenvironmental situation.

There must also be support for continuing research onthe plants that are currently listed or are candidates for

88

listing as endangered or threatened species. The publicrequires clear demonstration that a plant is indeedthreatened or endangered.

Also, the federal government needs to work closelywith commercial nurseries in facilitating the propagationof and trade in rare and endangered succulents. Theavailability of propagated plants will almost certainlyreduce the pressures from collectors on plants in the wild.

MexicoW. A. Fitz Maurice and Edward F. Anderson

Succulent plant floraThe Republic of Mexico has a great diversity of habitatsand flora. Succulents are found throughout the country,from the tropical forests in the south and south-east to thearid regions along the international border of mainlandand Baja California with the United States to the north.Two main mountain chains run down through themainland of the Republic: the Sierra Madre Occidentalin the west through the states of Sonora, Chihuahua,Durango, Sinaloa and Nayarit; and the Sierra MadreOriental to the east through the states of Puebla, Hidalgo,

Veracruz, San Luis Potosi, Nuevo Lebn, and Tamaulipas.Below this is the Trans-Mexican volcanic belt, and in theregion between these three ranges the high plateau ofnorth-central Mexico. All of these broad geographicalzones have a rich succulent flora, as do the arid and semi-arid zones of Puebla and Oaxaca, and the Baja Californiapeninsula. Although less diverse, there is also a uniquesucculent plant component in the vegetation of the low,hot, humid coastal regions and the southern and south-eastern tropical forests.

The extensive desert zones of Mexico consist of theChihuahuan Desert, including parts of the states ofChihuahua, Coahuila, Durango, Zacatecas, San LuisPotosi, Nuevo Le6n, Tamaulipas, Guanajuato, Quer@taro,and Hidalgo; the Sonoran Desert of Sonora and northernBaja California; and the Vizcaino Desert, including partsof the states of Baja California and Baja California Sur.

The main vegetation zones in which succulent groupsare found are the Chihuahuan Desert, Sonoran Desert,Vizcaino-Magdalena Desert, Desert Shrubland, TropicalShrubland, Tropical Forests, and the Oak-Pine Forest.However of the 32 types of vegetation in Mexico(Rzedowski 1986) all but the aquatic, semiaquatic, andp5ramo types contain succulents. The Matorral Xercifiloand its subdivisions Drobablv have the greatest diversitv of

Box 3.5 Estimated number of succulent genera and species per family in Mexico

Cactaceae: There is some disagreement in the number of taxa of Mexico in this family. Bravo-Hollis and Shnchez-Mejorada(1978, 1991) recognise 1080 taxa (854 species and 226 varieties and forms), whereas Hunt (1992) lists 925 provisional and fullyaccepted species. According to HernAndez and Godinkz (1994), following Hunt’s taxonomic framework, there are 563 wellrecognised species. Cactus nomenclature here follows Hunt, with the addition of new discoveries and published varieties.

Crassulaceae: Following Walther (1972), together with recent discoveries, there are over 150 taxa of Mexican Echeveria.Jacobsen (1960) lists 29 Mexican taxa of Dudleya, more than 30 of Villa& and 89 of Se&m. Additionally, using Jacobsen andincluding recent discoveries, there are 13 species of Pachyphylum, 14 of Graptopefalum, 4 of Lenophyllum, 3 of Thompsonella, 1of Pa&us and 3 of Kalanchoe. Thus, there are of the order of 350 Mexican taxa of Crassulaceae.

Agavaceae: Gentry (1982) recognises over 140 taxa of Agave. In addition, there are approximately 21 species of Manfreda(Pifia-Luj&n 1978) together with 11 of Furcraea, 3 of Hesperaloe, and 23 of Yucca (Standley 1920-1926 and Jacobsen 1960). Thisgives a total on the order of 200 taxa for the family in Mexico.

Fouquieraceae Henrickson (1969) lists 12 species of the single genus in this family, Fouquieria, all of which are found inMexico.

Other families: Other families with succulent representatives in Mexico, so-called caudiciform succulents, or speciesconsidered ‘succulent’ by collectors, include:

Aizoaceae: Carpobrotus (1) and Mesembryanthemum (1) . Anacardiaceae: Pachycormus (1) . Apocynaceae: Plumeria (5).Asclepiadaceae: Gonolobium (several), Asteraceae: Senecio (at least 2). Begoniaceae: Begonia (several). Bombacaceae:Ceiba (4) and Bombax (2). BromeHaceae: Hechfia (several). Burseraceae: Bursera (2 or 3) and Beiselia (1).Commelinaceae: 77adesccanfia (several), Convolvulaceae: lpomoea (at least 3). Cucurbitaceae: IbewNea (at least 2).Dioscoreaceae: Tesfudinaria (1). Euphorbiaceae: Euphorbia (at least 4), &tropha (several) and Pedilanthus (several).Fabaceae: Erythrina (several). Lentibulariaceae: Pinguicula (several). Li I iaceae: Aloe (1) . Moraceae: Dorstenia (several)and Ficus (at least 3). Nolinaceae: Beaucamea (6), Dasylirion (15), Nolina and Cal&anus (18). Oxalidaceae: Oxak (several).Piperaceae: Peperomia (at least 1). Portulacaceae: Portulaca (several) and Talinum (several). Urticaceae: Pilea (1) .Vitaceae: Cissus (at least I).

Totals: Overall, aside from the cacti and excluding orchids, this gives a minimum of 750 succulent taxa, and, including plantsas yet unnamed, there are possibly as many as 1000. Thus, there is probably a total of between 1600 and 2000 Mexican cactiand other succulent plants. The total number varies according to the definition of ‘succulent’ followed.

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Table 3.14 Main vegetation types and cactus diversity (Source: A&S my

Vegetation type Extent of land coverage(approx.) percent

No. of genera Percent of genera

Xerophytic scrub 45 39 34Tropical deciduous forests 17 24 21Conifer forests 21 7 6Tropical perennial forests 11 6 5Moist montane forests 1 4 3

Table 3.15 Uses of succulents in Mexico

Family Main uses

AgavaceaeCactaceaeCrassulaceaeFouquieriaceae

food, fibre, beverages, ornamentals, animal food, medicine, ceremonyfood, fodder, fencing, ornamentals, medicine, ceremonyornamentals, medicinefencing, ornamentals

succulents, particularly of the families Cactaceae,Agavaceae and Fouquieriaceae. (Table 3.14)

There are at least 28 plant families in Mexico whichinclude taxa that could be described as succulent (Box3.5). Of these, the four most important are the Cactaceae,Crassulaceae, Agavaceae, and Fouquieriaceae.

Local uses

Many succulent plants are utilised by Mexicans, assummarised in Table 3.15, and commercialisation ofvarious species is an important economic factor, especiallyin northern Mexico. The utility of succulent species canprovide an incentive for their propagation andconservation. There are, however, some instances ofoverexploitation of wild populations for local use asshown in the section on threats below.

The genus Opuntia (Cactaceae) is widely cultivated forits fruits (tunas), both in commercial orchards and onsubsistence farms. In addition, Opuntia (Platyopuntia)fruits are extensively collected from wild plants for salealong roads or in local markets. These species are alsogrown for ‘nopales’ (the new stems or ‘pads’ are skinnedand sliced to provide the basic green vegetable forsubsistence farms) and for cattle feed. The fruits of othercactus genera, including Escontria, Myrtillocactus,Polaskia, Stenocereus, and Hylocereus, are also harvested.

Various species of cacti are in demand for theirtraditional medicinal use. Mammillaria limonensis, forexample, is acclaimed in its habitat area as a remedy forsore throats; Opuntia jUgida is used to make a treatmentfor diarrhoea and other species of Opuntia have beenused as a treatment for diabetes; Pachycereus pecten-aboriginum and Lophophora spp. are used to treatrheumatism and other inflammatory ailments; andAriocarpus kotschoubeyanus, and other species, as a painkiller.

The genus Agave (Agavaceae) is of considerablecommercial value both for the production of alcoholic

beverages (mescal, pulque, tequila) and as a source offibres (istle, sisal, henequen). The sisal industry, decliningin the face of competition from plastics, usually involvescommercially grown plants and does not destroy theplant. Likewise Agave tequiluna, produced for tequila inSonora, Nayarit, and Jalisco, is derived exclusively fromcultivated stands.

Cuttings from Fouquieria species and Stenocereusmarginatus are widely used as barriers, often becomingliving fences. The latter is so widely used throughoutMexico that its geographic origin is uncertain. Species ofAgave are also used as living barriers. Cuttings fromOpuntia (Cylindropuntia) species are often used on thetop of low stone walls to make existing barriers moreformidable.

Threats

Habitat modification and conversion - Generally, thegreatest threat to succulent plant populations is changingland use. This is a side effect of the long term rapidgrowth of the Mexican economy, which has also resulted

Stenocereus fence, San Luis Potisi, Mexico.

90

in population migration to cities to meet the needs ofexpanding industries.

Amongst the numerous factors involved in thedisturbance of succulent plant populations, the conversionof land for both commercial and subsistence agriculturalpurposes is the most significant. For example, large areasof the succulent-rich Valley of Jaumave are beingconverted to agriculture, including the commercialgrowing of Aloe Vera. In other areas of northern andcentral Mexico succulent habitat is converted to maizecultivation as in San Luis Potosi where Ariocarpuskotschoubeyanus has become threatened. Agriculture isbeing increasingly practised on marginal desert land. Acommon practice throughout Mexico is the burning ofgrasses and low brush at the height of the dry season, inthe belief that nutrients are thus being returned to thesoil, and to avoid a build-up of tinder that could result in amore devastating fire. Many cacti and other succulentplants are lost or badly damaged in these brush fires, andthis practice of periodic burning of grazing areas does notappear to be controllable.

Land development for livestock production followsthe demands of the growing economy. While the majorimpact is from commercial use, there is a substantialimpact on the land from the goats, sheep and, to a lesserextent, cattle kept by the rural population. Plants aredestroyed in land preparation, and by trampling andgrazing by animals, as well as by chemical changes in thesoil. The resulting erosion causes further damage.

Destruction of habitat is also caused by road building,mining, the construction of dams, expansion of urbanareas, and industrial development, all of which lead to theloss of succulent plant populations. The new dam in theRio Moctezuma Valley has destroyed most of theremaining habitat of Echinocactus grusonii. Considerablehabitat destruction is also occurring in Baja California asa result of all-terrain vehicles. This, in addition toexpansion of irrigated agriculture and tourismdevelopments on the Magdalena Plain and the Caperegion, has led to the loss of some critical succulent plantsites.

In the Valley of Tehuacan-Cuicatlan between thestates of Puebla and Oaxaca, the construction of a newhighway from Mexico City to Oaxaca has destroyed partof the habitat of Cephalocereus hoppenstedtii, Agavetitanota, and Fouquieria purpusii. The habitat ofAriocarpus agavoides near Tula, Tamaulipas, is threatenedby the expansion of an urban garbage dump. UNAMperformed a rescue operation prior to the construction ofthis new super-highway.

Indirect, and less easy to define, universal problemsconcomitant to economic growth also threaten succulentplant populations. These problems include industrial,residential and vehicular pollution, diversion of naturalwater, and soil erosion.

1ZZega1 plant coLlection - Amateur and commercial

Marcos Sierra, horticulturist with Can Te, A.C,beside fouquieria purpusii at the edge of the newtoll road.

collecting of cacti and other succulents has been OIIC of

the major threats to wild species in Mexico, and remains asignificant problem. The commercial exploitation ofcactus populations in particular has led to the virtualextinction of several local populations of plants. Examplesinclude the tact i Turbinicarpus view&ii, Pelecyphorastrobiliformis, Mammillaria pectinifcra, Lophophoradiflusa, and Astrophytum asterias. Repeated collecting atfairly well-known sites in Mexico led to the neardisappearance of several of these species. There is someevidence that the seed reservoirs, given sufficient timeand protection, may allow the populations to return. Thismay be occurring, for example, at one of a number of sitesof Pelecyphora strobiliformis that had all the adult andjuvenile plants removed by commercial collectors in the1960s.

The market in the USA for illegally-collected Mexicancacti has declined recently as a result of stricterenforcement of legal controls at the Mexico-USA frontierand within the USA. Collecting for commercial gain bycentral Europeans remains a significant threat however.As an example, following the publication of the

91

Part of the Can Te, A.C. crew participating in a twoweek rescue operation with rising waters ofAguamilpa dam in Nayarit.

rediscovery of Mammillaria schwas-zii, dedicated collectorsbacktracked the deliberately vague site location data andfound the plant at its only known habitat. Thisinformation was then distributed among collectors withthe result that a population of around 1000 plants hasbeen decimated to about 100 individuals in five years,entirely due to collecting. This has happened despite thefact that artifically propagated plants of the species havebeen widely and legally available.

Although Mexican laws, international treaties, andincreased international border cooperation are reducingthe collection and illicit export problems, action on thepart of international conservation organisations is stillneeded to help control all aspects of the trade.

The problem of overcollection is aggravated at somerare plant sites, for example, those of Ariocarpusagavoides, Astrophytum asterias, and Mammillan’a hererae,where local people realised the financial rewards to begained by selling plants to foreigners. Collected plants areremoved from their habitat in quantities well in excess ofdemand, and are poorly cared for. Similarly, the demandfor rare plants has given rise to local guides well known tothe foreign collecting community.

Collection of cacti and succulents for use withinMexico also threatens wild populations of some species.Small succulent plants are collected and potted and arewidely available in Mexican nurseries for decorativepurposes. Large cacti such as Ferocactus are sold asgarden plants. In rural Mexico, as well as in some cities, itis the custom to make up a nativity scene or ‘nacimiento’each Christmas. Traditionally, this includes smallMammillaria species such as M. plumosa, M. bocasana, M.albicoma, M. dumetorum, and M. aureilanata. These plantsare collected commercially and shipped to marketsthroughout Mexico. Often the children in rural areas aresent into the hills to collect suitable plants for their own

family’s use. The plants are generally discarded after theholidays although they are sometimes cultivated.

Although agaves are commercially grown on a largescale to supply distilleries, many small mescal and pulqueoperations use agaves gathered from the wild, and this isresulting in local depletions. Mescal production in Sonorahas markedly reduced the number of native agavesreaching flowering age. As a consequence, the bats, whichare the chief pollinators of those agaves and manyimportant cacti in the area, have become significantlythreatened.

Species of Ferocactus, Melocactus, Echinocactus, andother similar cactus forms are widely collected for smalloperations making ‘crystallised fruit’ (the plant bodies arecut into cubes and boiled in syrup). A group of producers

g in the Queretaro area recently approached the5 organisation Can Te, A.C. asking for assistance in large

scale propagation of Ferocactus histrix, as this principalsource of material for their industry was becomingincreasingly difficult to find in habitat.

Species of barrel cacti, for example Echinocactusplatycanthus and Ferocactus pilosus, are used extensivelyin northern Mexico as forage and as a water source forgoats and, less frequently, cattle especially during the dryseason. This practice along with other uses of the plantshas resulted in a dramatic decline of the populationdensities of these species. The same has happened withwild Opuntia (Platyopuntia) collected for cattle feed innorthern Mexico.

The use of Lophophora williamsii or ‘peyote’ is wellknown. The species is listed in Mexico as a narcotic withmulti-year prison terms awarded for possession. Its statusas a narcotic has given rise to destruction of both L .williamsii and other, often rarer species of similarappearance by zealous officials. Educational materialsdealing with identification of the species involved arecurrently in preparation for widespread distribution tolocal authorities. The government allows the collecting

Lophophora williamsii, the true ‘peyote’, in flower inhabitat, San Luis Potosi.

92

* SEDUE (Secretary of Urban Development and Ecology) wassuperceded in the area of ecology, in the early 1990s bySEDESOL (Secretary of Social Development). Under the newPresident, Ernest0 Zedillo, SEDESOL was further refined toSEMARNAP (Secretary of the Environment, Natural Resourcesand Fish). Within that organisation, INE (National Institute ofEcology) is the institution most actively involved with threatenedspecies and collecting permits.

W

Drying peyote, Texas.

and useindigen0

in the religious ceremonies of someof peyoteus peoples.

Conservation status

The impact of all the various threats on Mexican cacti ando ther succu len t s depends on the popu la t ioncharacteristics of individual species. The majority ofMexican endangered cacti occur in small, disjunctpopulations, primarily in arid and semi-arid regions of thecountry, and a significant proportion of them arerepresented only by one or a few populations. Most ofthese species have a combination of biological andecological attributes making them extremely vulnerable toany form of disturbance. These plants usually have slowgrowth rates, long life cycles, and the recruitment of newindividuals in the population is extremely low. Theseinherent characteristics, along with the peculiarbiogeographical patterns of these plants, determine a slowdemographic response of the populations afterdisturbance. Unfortunately, accurate plant populationinformation is generally not available and so it is oftendifficult to evaluate the precise degree of threat toindividual species.

Various preliminary studies of the conservation statusof Mexican succulents have been carried out over the pastfifteen years. The Threatened Plants Committee (TPC) ofIUCN undertook a survey of the conservation status ofMexican Cactaceae in the early 1980s mainly bycorrespondence with experts in Mexico and elsewhere.IUCN categories of threat were applied to all Mexicanspecies and this formed the basis for the data on Mexicancacti held by WCMC. Data holdings at WCMC haverecently been revised by harmonising with the CITES

Cactaceae Checklist (Hunt 1992) and information onthreatened Mexican cacti and other succulents publishedby SEDESOL and subsequently SEMARNAP”.

Concern about the situation in the wild of MexicanCITES Appendix I cacti led to the field survey workconducted by members of the SSC Group and Mexicanconservation trainees from 1986 to 1988, funded byWWF-US. The results of this field work (Sanchez-Mejorada et al. 1986; Anderson 1990; and Anderson et al.1994) confirmed the conservation status of a range of 52rare and threatened species.

More recently, a newly formed group working at theInstitute of Biology, UNAM, has centred its researchactivities on the study of the biogeographical patterns ofthe Mexican endangered species, particularly thosegrowing in the Chihuahuan Desert Region (Hernandezand Godinez 1994; Hernandez and Barcenas 1994). Overthe past few years this same group has assembled adatabase of herbarium collections from North andCentral American Cactaceae containing so far(September 1994) over 9500 records from 37 Mexican,USA, and European herbaria (Hernandez et al. 1993).This is the largest available database containinggeographical data on cacti species, and is provinginvaluable to determine the areas of species concentrationof endangered cacti in Mexico.

It has been estimated (Hernandez and Godinez 1994)that 73 per cent of the genera and 78 per cent of thespecies of cacti occurring in Mexico are endemic, and that35 per cent (197 species) are somewhat endangered. Inabsolute terms, the country possesses the highest numbers

Mexican cactus and plant collector with Pelecyphorastrobilifonm’s, Tamaulipas, Mexico.

93

Pachycereus pringlei, Desert Botanical Garden,Arizona.

of endemic and endangered cacti, comparing with othercountries such as Chile, Ecuador, Cuba, and Brazil whichalso have highly significant proportions of endemic andendangered species.

Currently field surveys to assess the status of Mexicancacti are being undertaken by scientists from variousbotanical gardens and universities within Mexico.Botanists at UNAM are, for example, working onecological and population studies in the ChihuahuanDesert area in the Tehuacan-Cuicatlan Valley and havebeen studying the conservation status of cacti in the Stateof Queretaro for over ten years. At a recent conferenceon population studies held at Can Te, A.C. in San Miguelde Allende, Guanajuato presentations were given on workin progress by researchers at the Universidad Autonomade Tamaulipas, the Centro de Investigaciones Biologicasde Baja California Sur and the ITESM, CampusQueretaro. Can Te, A.C. is continuing to work with theseinstitutions and others to pull together the diverseinformation available. A comprehensive program ofdetailed population studies is urgently needed to build onthe work already in progress. It is clear that this must becoordinated and implemented locally.

Few detailed studies have been made of the long-term

fluctuations of cacti and other succulent plant populationsin habitat. These are necessary to understand the naturalbehaviour of populations and the reasons for their rarity,and, via comparative studies, to assess the basis of theirvulnerability. Can Te, A.C. started such investigations in1991 and currently has 50 ongoing studies with others inpreparation. Concomitant with further population studies,additional long term studies can and need to be initiated.

A CITES-financed project is currently in progress tocarry out population studies on Mexican cacti and toevaluate the impact of legal and illegal trade on wildpopulations of these species. The work is being carriedout by Can Te, A.C. and the Desert Botanical Garden,Phoenix, with the assistance of several local botanicinstitutions.

Information on the conservation status of MexicanAgavaceae is included in Annex 1. A list of threatenedMexican cacti and other succulents as prepared bySEMARNAP is given in Annex 11. The cactusinformation within that list is based on Hernandez andGodinez (1994) which modified Hunt’s (1992) CITESCactaceae Checklist.


The Chihuahuan Desert Region, which is the largest and

5least understood North American Desert, contains the

2 largest assemblage of endangered cacti in the wholeg continent. Hernandez and Barcenas (1995) have2li evaluated the major areas of concentration in this regionti in terms of their species richness along with their

abundance of rare, geographically restricted cacti. Withinthe Chihuahuan Desert the species are aggregated inareas of moderate altitude, particularly towards the south-eastern, and to a lesser extent, the eastern margins of thisdesert, in northern San Luis Potosi, and in southernCoahuila (including the Bolson of Cuatro Cienegas),Nuevo Leon and Tamaulipas. Another area of similarimportance is the Queretaro-Hidalgo Arid Zone whichincludes areas such as the Rio Extorax Basin and theMetztitlan Valley.

The richest area within the Chihuahuan Desert islocated in northern San Luis Potosi, around a localityknown as Huizache, where a total of 14 endangeredspecies are recorded. Towards the north and north-east ofHuizache there are several other areas where animportant number of endangered species occur. Thesecorrespond to regions near Matehuala, Doctor Arroyo,Galeana, Aramberri, Jaumave, and Cuatro Cienegas.Another area of similar importance as the Huizache is theRio Extorax Basin, in the State of Queretaro. This areawhich is separated from the main body of the ChihuahuanDesert, corresponds to the Queretaro Arid Zone and issurrounded by several cactus-rich regions located inGuanajuato, Queretaro, and Hidalgo.

Major areas of succulent diversity in Mexico includethe following:

94

1)

2)

3)

4)

5)

6)

Chihuahuan Desert Region, including disjunctportions in the Jaumave and Tula valleys, and theQueretaro-Hidalgo Arid Zone. According to thestudies of Hernandez and Barcenas the areas withinthis region that merit inclusion within Mexico’sNational System of Protected Areas are:

a) Toliman (Extorax Basin, Queretaro),b) Huizache (San Luis Potosi),c) Metztitlan, (Hidalgo),d) The valleys of Tula and Jaumave, Tamaulipas -

these have a high percentage of rare andthreatened cacti which are very popular withforeign collectors, including Ariocarpus agavoides,Turbinicarpus saueri, Tysabelae, Obregonia denegrii,Pelecyphora strobiliformis, and Mammillaria zubleri.

Another site in this region identified as beingimportant for conservation is Doctor Arroyo north toGaleana and Rayones, including Aramberri Valley, inNuevo Leon State.

Sonoran Desert including the Cape Region of BajaCalifornia Sur, the islands of the Gulf of BajaCalifornia together with the Pacific Ocean West Coastof Baja California. The area around Catavina in thesouthern part of Baja California State is especially richwith Fouquieria (Idria) columnaris, Pachycormusdiscolor, and other important species such asFerocactus gracilis, Pachycereus pringlei, Washingtoniarobusta, and Dudleya spp.

Tehuac6nXuicatl5n Valley, Puebla-Oaxaca: densewooded sites of arborescent cacti with Cephalocereushoppenstedtii, Escontria chiotilla, Ferocactusflavovirens, F. haematacanthus, F. robustus, Mitrocereusfulviceps, Myrtillocactus geometrizans, Neobuxbaumiamacrocephala, N. mezcalaensis, N. tetetzo, Pachycereushollianus, P. weberi, Pilosocereus chrysacanthus, as wellas Mammillaria napina, M. pectinifera, and othersucculents such as Agave spp., Fouquieria purpusii, andBeaucarnea gracilis. UNAM has studied the area indetail and proposes that the least disturbed area,south of the city of Tehuacan, should be protected.

La Mixteca Alta (Oaxaca)

Balsas Basin (Oaxaca , Puebla, Guerrero, andMichoacan)

Isthmus of Tehuhntepec (Oaxaca)

Other succulent rich sites of outstanding importance are:

7) Barranca de MetztitlBn, Hidalgo: spectacular habitatfor unique “viejitos” Cephalocereus senilis, withAstrophytum ornatum, Echinocactus platyacanthus,Fouquieria fasciculata, Mammillaria geminispina, M.humboldtii.

8) Habitat of Aztekium hintonii and Geohintoniamexicana with Mammilloydia candida var. caespitosa,M. picta, M. winterae, Thelocactus tulensis var.matudae, Yucca sp. cf. carnerosana, and others.

9) Pedregal de Cuernavaca (Chichinautzin), Morelos:one of the most beautiful natural succulent gardens inthe Mexican Republic with Agave horridu, Echevcriagibbiflora, Sedum frutescens, S. oxypetalum, manyorchids, begonias, and bromeliads.

These areas, rich in succulent taxa and of outstandingbeauty and interest, merit inclusion within the NationalSystem of Protected Natural Areas. In addition, thefollowing succulent-rich areas have also been identified ascritical for the protection of their native flora:

a>b)4d)e>f >g>h)0.J>W

El Canon de1 Zopilote, Guerrero StateThe Desierto de Altar, Sonora StateViesca-Saltillo area, Coahuila StatePiedras Negras-Sabinas area, Coahuila StateMapimi region between Durango, Chihuahua andCoahuila StatesBolson de Cuatrocienegas, Coahuila StateCumbres de Monterrey, Nuevo Leon StateBarranca de Tolantongo, Hidalgo StateMesa de1 Nayar, Nayarit StateHuayacocotla, Veracruz StateLlanura de1 Rio Verde, San Luis Potosi State


Mexico has made major progress in succulent plantconservation activities in the past decade. Existing lawsare being effectively enforced, and scientists are activelyengaged in research related to conservation. The effectsof these activities are already noticeable on wildpopulations of succulents. Illegal collecting and thewanton destruction of plants have been greatly reduced.

The decline of many succulent plant populationscontinues, however, and sustained efforts on a broad frontwill be necessary to prevent further extinctions. As far aspossible, local inhabitants need to be involved inconservation planning to ensure full support forconservation decisions.

Protected areasThe Mexican National System of Protected NaturalAreas, under the direction of various ministries, includes73 protected areas. These areas fall into nine categories:Biosphere Reserve, Special Biosphere Reserve, NationalPark, Natural Monument, Aquatic Park, Area ofProtected Natural Resources, Area of Protected WildFlora and Fauna, City Park, and Ecological ProtectionZone. A total of over six million hectares or 60,000 km’are currently under protection. About half of the totalarea covers arid land, but relatively few protected regionsexist in the northern part of Mexico where there is thegreatest diversity of succulents.

95

The existing protected areas on paper are generallyneither staffed nor fenced on the ground, but they offerthe potential to provide special protection where a highthreat category has been identified. Greater financialsupport for the existing federal protected areas is needed.Protected areas with numerous succulents include:

a>b)44e>9g>h)9.J>k)

El Pinacate ReserveMapimi Biosphere ReserveCanon de la HuastecaParque International de1 Rio BravoIsla Cedros SanctuarySierra de Manantlan Biosphere ReserveIsla Isabel National ParkRio Lagartos Ecologic ReserveCanon de1 Sumidero National ParkLagunas de Chacagua National ParkSan Pedro Martir National Park

In addition, a large region near Cuatro Cienegas has beenproposed as a protected area.

The habitats of threatened species, Cremnophilanutans (Crassulaceae) and the yellow-spined variant ofMammillaria spinosissima fall within the confines of theTepozteco National Park in the state of Morelos.Ferocactus tiburonensis is endemic to the island ofTiburon and the entire island is a special reserve. The bestknown habitat of Agave victoriae-reginae lies within thearea of Cumbres de Monterrey, the largest national parkin Mexico.

At present La Comision National de la Biodiversidad(CONABIO) p pis re aring a list of the additional areas inMexico appropriate for designation as protected areas.Several areas with populations of succulent plants need tobe protected, provided such action does not create heavyeconomic and social burdens on the people living in thevicinity. Local inhabitants, if included in decisionsinvolving regions near their villages, may provide thegreatest security for the plants. Perhaps financialincentives might enable them to steward these areas aswardens or guards.

Legal protection for succulentsThe General Law of Ecological Balance and Protection ofthe Environment, passed by Mexico’s Congress in 1986,establishes the principles and regulations by which allspecies of wild flora should be treated and used inMexico. All uses of wild flora are covered by this law anrequire authorisation of the Secretary of Environmen tNatural Resources and Fish (Secretaria de MediAmbient, Recursos Naturales y Peces or SEMARNAP >specifically of the National Institute of Ecology (Institut .cNational de Ecologia or INE) through its Gener;Administration of Ecological Use of Natural Resources(Direction General de Aprovechamiento de 10s RecursosNaturales or DGAREN). Permits for the collection of anywild plants are required by the Mexican government,which the head of DGAREN must personally sign.

Infractions of the law range from misdemeanors tofelonies. It is the responsibility of SEMARNAP to bringsuit in the case of misdemeanors and to refer suspectedfelonies to the Federal Attorney General (ProcuraduriaGeneral de la Republica or PGR). The law also allows anindividual to bring suit where it is believed the law hasbeen violated.

Another law which has relevance to succulent plantconservation is the Federal Forestry Law (Ley FederalForestal). This law was recently amended (late 1996) andnow, once again, includes and regulates non-woodyplants. Perhaps even more significant is the typification inthe Penal Code of illegal removal of plants, includingcacti and other succulents, making such illegal removal a‘delito ambiental’ (an environmental crime) with jailsentences now possible of three to six years.

Existing legislation is sufficient to protect the nativeflora and fauna of Mexico, but greater efforts are neededto educate the public about federal regulations.SEMARNAP is required to list the species of flora andfauna to be regulated (Annex 11). In addition, violators ofthe law should be prosecuted to emphasise that theMexican government is serious about preserving itsnatural wealth. Table 3.16 provides examples of someenforcement acts.

Pachycereus pecten-aboriginum, a cactus of Mexicoused in local medicine.

96

CITESMexico is a recent signatory of CITES, and SEMARNAP,as the Managment Authority, has the responsibility ofenforcing regulations pertaining to the international tradein succulents.

At present there appear to be misunderstandings bymany people about the purpose of CITES. ManyMexicans and foreigners do not understand the permitsystem presently in operation by SEMARNAP, and nearlyall foreign collectors have ignored the system. In addition,inspectors at the international borders need training inthe identification of plants currently controlled byMexican law.

More effective and timely means must be found forrepatriating Mexican plants confiscated at foreign bordersor seized in foreign countries. This will require puttingpressure on the authorities in the countries where theplants are confiscated to assign responsibility forrepatriation costs.

Ex situ conservationMexico has 42 registered botanic gardens listed bySEMARNAP (Annex 13). Coordination of theconservation activities of Mexican Botanic Gardens isfacilitated by the Asociacion Mexicana de JardinesBotanicos. The Asociacion is promoting the formation ofnational collections for various groups of plants. Thenational collection for Agavaceae is already maintainedby the Jardin Botanic0 de1 Instituto de Biologia, UNAM.The collection currently holds 80 per cent of the Mexicanspecies in the family.

Botanic Gardens are playing an important role in‘rescuing’ succulent plants from sites threatened bydevelopment projects. In 1990, UNAM conducted anintensive rescue operation in the Rio Moctezuma valley,the habitat of Echinocactus grusonii, as well as other rareplants, prior to the major dam project. In 1992, Can Te,A.C. conducted a rescue effort at a well known site ofPelecyphora aselliformis and Mammillaria aureilanatathreatened by construction of a highway. At the request ofthe Mexican government in 1993, plants were rescuedfrom a 10,000 ha area about to be flooded by a new damat Aguamilpa, Nayarit. This rescue operation wasundertaken by Instituto de Botanica de la Universidad deGuadalajara, and La Escuela de Biologia de laUniversidad de Guadalajara, together with Can Te, A.C.

Site of rescue operation with Pelecyphoraaselliformis and Akmmillaria aureilanata. From left:Gabriel Solano of SEMARNAP in Mexico City; Biol.Emilio Baltazar Cuellar Jimenez from the San LuisPotosi office of SEDUE; W. A. and Betty FitzMaurice, investigators with Can Te, A. C.; andCharles Glass, Curator of Plants for Can Te, A.C.

In another important rescue operation UNAM relocatedspecimens of around 50 species from the site affected bythe construction of the Tehuacan-Oaxaca highway. Theplants were placed in the Botanic Gardens of UNAM, theCentro Interdisciplinario de Investigation para clDesarrollo Integral Regional-Unidad Oaxaca (CIIDOR-Oaxaca), and African Safari Parque Zoologico enValsequillo, Puebla.

In addition to caring for and propagating the rescuedplants in botanic gardens in Mexico, consideration is alsobeing given to reintroduction of plants to suitablehabitats, as UNAM has done at the Rio Moctuzema site.For example, UNAM’s tissue culture work has allowedthe reintroduction of Mammillaria sa12-a~~gelcllsis inMexico City.

Botanic gardens also act as rescue centrcs for cactiand other succulent plants confiscated from collectors atvarious frontiers and occasionally within Mexico. Seizedplants are sometimes sent to Mexican institutions, andcollections are being maintained, for example, by Jar-dinBotanic0 de1 Instituto de Biologia, UNAM; Instituto de

Table 3.16 Enforcement actions by Mexican authorities 1991 and 1992

People prosecuted Plants collected Penalty

4 Austrians 878 Appendix I and II specimens4 Belgians 1569 Appendix I and II specimensI German Appendix I and Appendix II cacti1 German 423 Appendix I and II specimens3 Italians c. 400 Appendix I and II specimens1 American 28,624 Appendix I and II specimens

each fined US$1200each fined US$800prison sentenceUS$425each fined US$400returned to USA before the case was decided

in court

97

Part of a plantation at the Botanic Garden, El Charcodel Ingenio, San Miguel de Allende, of 500 goldenbarrels (Echinocactus grusonii) whose naturalhabitat is currently under the waters of the newZimapan dam on the border between Quektaro andHidalgo.

Botanica, Universidad de Guadalajara; UniversidadAutonoma Agraria Antonio Narra; and Can Te, A.C., butthis has so far been on a somewhat random basis. Thesheer number of plants involved can place a strain onexisting resources of the botanic gardens.

Nursery developmentIn Mexico there is extensive nursery production of plantsfor the domestic market, but little of this has beendevoted to cacti and other succulent plants. Until recentlythere were only two commercial nurseries propagatingsucculents from seed and cuttings. The situation is nowchanging with a rapidly increasing consumer interest innative succulents. This coincides with intensified effortsby the Mexican authorities to control the trade in wild-collected plants. Sixteen nurseries have made applicationto the authorities for permission to propagate succulentplants (see Annex 12). Although still small, the industryshows good growth potential.

Main agencies involved in conservationThe main governmental agency responsible forconservation in Mexico is the Consejo National de

Ciencias y Technologia (CONACYT). Both SEMARNAPand the Secretary of Agriculture (Secretaria deAgricultura y Recursos Hidraulicos or SARH) areresponsible for the control of the use of Mexico’s naturalflora; the direct concern lies with SEMARNAP. Manyother governmental agencies are directly and indirectlyinvolved.

In addition to the governmental agencies a number ofuniversities and many non-governmental organisationsare active in conservation efforts relating to succulentplants. These include:

Departamento de Botcinico, Instituto de Biologia de laUniversidad National Authoma de M&co (UNAM) housesthe National Herbarium of Mexico, which has the largestcollection of Mexican plants (over 550,000 specimens),including an important set of herbarium specimens ofCactaceae, Agavaceae, Crassulaceae, and several othersucculent plant families. A research group at thisinstitution is devoted to generate basic information on thetaxonomy, biogeography, and ecology of rare andendangered cacti, particularly from the ChihuahuanDesert Region. Also this group has developed the largestdatabase of herbarium collections of North and CentralAmerica, currently containing over 9500 records from 33institutional herbaria.

Jardin Bothico de1 Znstituto de Biologia de la UniversidadNational Authoma de Mbxico (UNAM) in Mexico City hasa well-established and active conservation programmerelating to the study and conservation of cacti and othersucculents. Activities include:

study of the cactus flora of Meso-America, includingfield collections for the National Herbarium;study of the biogeography, distribution andconservation status of the cacti of the TehuacanValley, and the development of a proposed protectedarea;publication of scientific materials on succulents;maintenance of the national collection for Agavaceae;propagation of endangered cacti including thedevelopment of tissue culture techniques;development of educational materials relating to cactiand agaves.

Can Te, A.C., a non-governmental, not-for-profitorganisation dedicated to conservation, has a newlyformed botanic garden that is taking a lead in succulentconservation work. Several important salvage operationshave been carried out, and rescued plants are now beingpropagated. Can Te, A.C. also provides information onpropagation to producers and collectors, for example, tothe crystallised fruit manufacturers mentioned above. CanTe, A.C. also has a program, ‘Comprar para Conservar’ or‘Purchase to Preserve’, encouraging people to contributefinancially toward the purchase of threatened habitats.

Yd

Universidad Authoma de Tamaulipas - botanists from thisuniversity have been carrying out detailed populationstudies on endangered cacti in the Jaumave area.

Sociedad Mexicana de Cactdceas, the main specialist societyfor succulent plants in Mexico. It is involved in thedissemination of information on Mexican cacti and othersucculents and their conservation.

Centro de Investigaciones Biolbgicas (CIB) in La Paz -carries out ecological studies of cacti in the BajaCalifornia area.

Universidad de Guadalajara, where studies are carried outon the cactus flendemic species.

ora of Jalisco and the propagation of

ITESM-Campus Querbtaro carries out studies on the cactiof Queretaro, undertakes propagation activities,maintains a small gene bank, and has an educationprogramme about cacti of the state.

The authors would like to acknowledge the following people forkindly providing additional information and review of this section:Salvador Arias, Helia Bravo H., Robert Bye, Federico Gama,Abisai Garcia Mendoza, Keith Gardner, Charles Glass, KennethHeil, Hector M. Hernandez , Wendy Hodgson, George Lindsay,Reid Moran, and Gary Nabhan.

The West IndiesAlbert0 Areces-Mallea

The Caribbean islands form an archipelago of over 1000islands of considerable range in size, altitude, soil typesand environmental niches. Exposed land surfaces cover adistance of 2700 km from Barbados on the east, to thewestern tip of Cuba, while the distance from Grenada in

the Lesser Antilles to the northern tip of the Bahamas is1900 km. The islands range in size from Cuba, with114,500 km2 and a vascular flora of over 6000 species, toislets of small rocks of a few square metres and a flora ofa dozen species.

The diversity of the West Indies is seen in such factorsas altitude of the islands, temperature range, soil types,and units of vegetation on each island. Hispaniola has thegreatest range in altitude where the Enriquillo Basin is 30m below sea level and Pica Duarte reaches 3000 m.Volcanic peaks in the Lesser Antilles range from 900 m toapproximately 1500 m. The major part of the land surfacewithin the archipelago is below 300 m in altitude.

The average (mean) temperatures of the area atelevations of less than 90 m range from 24.9” (Havana) to26.1 “C (Dominica). Neither annual nor daily variations intemperature are great. Day-length range is nearly twohours greater in Nassau in the north than it is in Trinidadjust south of the area under consideration. With respectto rainfall, many areas of the West Indies receive less than1000 mm of annual precipitation, while rainfall exceeding5000 mm has been suggested for several areas. Sometimesthere are six or seven months of reduced rainfalloccurring as two dry periods, while many mountain areasshow no months of rainfall of less than 100 mm.Succulents and low spiny shrub vegetation arecharacteristic of coastal areas with seasonally highertemperatures and rainfall under 700 mm, often falling inone short annual period.

The West Indies show a variety of soil types whichoffer a relatively large number of ecological niches. Thereare siliceous savannas in western Cuba and on the northcoast of Puerto Rico. The central portion of Cuba has anearly uninterrupted serpentine savanna. Areas ofgypsum and salt concentrations, often in bands, occur inHispaniola. A belt of aluminous lateritic soil extendsthrough most of the island of Jamaica and occurs also in

Cactus scrub withStenocereus hystrix andPilosocerus royenii withPlumeria obtusa in theforeground, MonaIsland, Puerto Rico.

the southern peninsula of Hispaniola; areas of volcanicactivity occur in the Lesser Antilles. Outcrops oflimestone as sedimentary rock or elevated coral reefs areabundant, and areas of intrusive igneous rock could beadded to the chart as additional specialised habitats. Thevegetation of the various soil types is often distinctive incomposition or habit and is frequently high in endemicspecies. In general, geologic and edaphic featurescombine with climate and topographic factors such ashigh relief and alternation of lowlands and mountains, todetermine the unique characteristics of the plantcommunities and the floristic diversity of each individual,isolated island.

The Caribbean Islands have a natural vegetationconsisting of lowland and montane tropical forest,evergreen thicket, savanna, cactus-thorn scrub, marsh orswamp, mangrove, beach, and riverine communities. Thetotal flora of the region consists of about 13,000 vascularplant species with around 6550 regional endemic species.About half of the endemics occur only in Cuba.

Institutional bases for the study of the West Indianflora locally are patchy and most of them are limited inphysical resources and capabilities. Fortunately, interest isgrowing and there is current activity in, for example,Cuba, Dominican Republic, Puerto Rico, St. Lucia, andBarbados. In recent years, Floras covering both floweringplants and ferns have been published for the Bahamas,Cayman Islands, Jamaica, and the Lesser Antilles. Aflowering plant Flora for Hispaniola is well advanced inproduction, although the Cactaceae has not yet beencovered. New Floras of Cuba and Puerto Rico are beingprepared. There is no current guide to the succulents ofthe West Indies.

A general Flora, to include all cryptogamic andphanerogamic groups, is proposed for the GreaterAntilles under the direction of The New York BotanicalGarden. Most of the current research on floristics in Cubaand Hispaniola is published in those islands, but for otherterritories, investigations and publication areinstitutionally based in North America and Europe.

The succulent floraThe succulent flora of the West Indies consists mainly ofplants in the Agavaceae, Apocynaceae, Cactaceae,Euphorbiaceae, Moraceae, Portulacaceae, and Vitaceae.Also included here are marginally succulent species suchas the West Indian Bombacaceae, a Sterculiaceae(Hildegardia) with a bulky, greenish trunk, all Dorstenia(Moraceae), and many of the native Euphorbiaceae withsomewhat thick stems or leaves more fleshy or thickerthan normal. However, no Burseraceae, Begoniaceae,Piperaceae, Rubiaceae, Urticaceae, aroids, or orchidsother than the succulent vinelike VaniZZa, well representedin the dry areas, are considered. Out of 324 so definedsucculent plant species recorded here for the WestIndies*, 243 (75 per cent) are endemic. Succulentscomprise about 2.5 per cent of the total number of

* In this account including the Florida cays (only their endemicspecies, not found in the mainland of Florida) but excludingAruba, Bonaire, Curac;ao, Trinidad, Tobago, Margarita, and 100

other small islands adjacent to Venezuela, which arephytogeographically part of northern South America.

flowering plants of this region. The remaining 81 species(25 per cent) extend to continental landmasses in North,Central, and South America, and even sometimes to theOld World tropics and/or subtropics. This latter group isnot of primary conservation concern. It is made up ofmany species occurring along the seashore, some withinbeach areas, coastal salt marshes, dunes or flats, andothers on maritime rock exposures.

The families containing succulent plants in the WestIndies are listed in Table 1 of Annex 14 with the numberof succulent species in each genus and an indication ofendemic taxa. No subspecific taxa are considered due tothe yet insufficient knowledge of the Caribbean flora.New succulent species continue to be discovered (Areces-Mallea 1992, 1993) and, despite the level of botanical andhorticultural interest in the group, the taxonomic status ofa number of West Indian succulents remains poorlyknown. The non-endemic succulent species native to theWest Indies are listed in Table 2 of Annex 14, whereas theendemics are shown in Table 3 of Annex 14 with theirgeographic distribution, regional or restricted, and anevaluation of their conservation status.

Stebbins (1952) pointed out that a dry environmentstimulates speciation. The ecological and chorologicalstudy of most of West Indian endemics provides evidencefor this phenomenon. In general the largest number andhighest density of endemics may be observed in the aridzones (coastal areas, semi-deserts) and in thephysiologically dry habitats (serpentines, limestone karsts,siliceous sands). Endemic succulent taxa are mostcommonly found within coastal plant communities.Cactus scrub, evergreen bushland and dry evergreenthicket occupy well drained, usually rocky, substrates.Relatively extensive semi-desert vegetation - uniqueflora of endemic cacti, other succulents, and spiny shrubsexist along the leeward coasts of the larger islands. Thevegetation types where most of the cacti and succulentslisted in Tables 1 and 3 of Annex 14 occur are describedhere.

I. Coastal formations1) sandy beaches - Herbaceous and shrubby vegetationof the tropical sandy sea shores, commonly distinguishedby two main associations: a) an open pioneeringcommunity formed of creeping lianes and stoloniferousgrasses (Cakile lanceolata and Blutaparon vermiculare areoften associates), and b) a less open community, the nextsuccessional stage, with Sesuvium portulacastrum,Chamaesyce mesembrianthemij’olia and Argusiagnaphallodes. On low sandy shores, seaside prairies, andsandy meadows Borrichia arborescens and Scaevolaplumieri might be abundant. Suriana maritima is moreoften found in littoral thickets in the transition of themeadows toward strand vegetation.

Sandy beach communities are common, though oflimited extent on the West Indian islands, when comparedwith the rocky or coralline coastal associations.

Undisturbed beach areas are increasingly difficult to find;most of the sandy shore areas have been heavily damagedor destroyed. Fortunately, most of the genera and speciesinvolved are of wide geographic distribution.

2) Strand Z&o& scrub and low forest - Inland from thecoastal beach, dunes may be built up consisting solely ofsand or of sand deposited on a rocky substratum. The sea-grape Cocc&ba uv$era is the classic component of suchstrand areas. Although the monodominancy of sea-grapestands is an essential characteristic found all over theWest Indian islands, there are also differences withrespect to the floristic composition of these associations indifferent locations. The community of sea-grape andOpuntia &ZZenii is relatively common on dry coastal areas,where the sand dunes are situated on shallow coral reefsor low banks. In the larger cays of northern Cuba Opuntiamillspaughii may also be present. Selenicereus spp. arecommon climbers on the sea-grape. Other associatedsucculent species found occasionally are Scaevolaplumieri, Suriana ma&ma, and Argugia gnaphallodes. Thestrand littoral scrub and low forest are still common onthe sandv shores and on the first seaward dunes of the lowlimestone rocky shores of the Antilles, Bahamas, andFlorida.

3) Sal ine f la ts - The vegetation of the salines iscomprised of leaf-succulent dwarf shrubs, annual orperennial succulents, and grasses of high osmotic tension.This belt of salt vegetation is developed on the inland sideof the mangrove-zone, in the areas flooded only twice ayear by the high equinoctial tides, where the salt tends toconcentrate by evaporation, and allows the developmentof herbaceous flats with Batis maritima, Salicornia spp.,and Suaeda spp. Other succulents occurring in the salineprairie vegetation, mostly at the edge of the supratidalbelt, are Heliotropium curassavicum and Blutaparonvermiculare. They are all of wide geographic distribution,and no endemic taxa occur in these communities. Due toits environmental requirements this saltwort vegetation isnot extensive, nor very abundant in the Caribbean islands.However patches and belts up to one kilometer wide maybe locally common in Cuba.

4) Rock pavement vegetation - This orophiloushalophytic vegetation of the supratidal rocky shores, isconditioned by the influence of salt spray, the unprotectedexposure and extremely poor soil conditions. The rock-pavement substratum for this type of vegetation occurs onuplifted coral reefs as benches. It dominates entire cays ofsuch origin or occurs in outcrops between sandy or lowmuddy beaches. There is a well-recognised pioneerassociation of creeping or prostrate leafsucculent plantssuch as Sesuvium spp., Trianthema portulacastrum andLithophila muscoides, and another association withBorrichia arborescens, Chamaesyce mesembrianthemifoliaand Opuntia diZZenii. This community that grows on the

inland side is characterised by more dense growth. InPuerto Rico and the Virgin Islands Opuntia repens may bealso present, and on Mona Island (Puerto Rico) there is aspectacular dominance of Mammillaria nivosa within thistype of vegetation. In contrast to sandy and salinevegetations, which are primarily of pan-tropical character,the vegetation of coastal rock pavements is mainly ofAntillean and Caribbean distribution.

II. Coastal-lowland formations1) Dry limestone shrubwoods - This dense vegetationcornposed of thorny, sclerophyllous, small-leaved treesand shrubs occurs on bare rocks of dry limestone terracesusually on the inland side of the coastal rock pavementvegetation, and on the lowland karstic ‘dogtooth’formations. The 2-3 m high shrubs and the emergentindividuals or groups of 5-6 m ‘rod-like’ associated treesare best developed under climatic conditions consisting oftwo dry seasons per year which together amount to aboutseven or eight dry months. Columnar or tree-shaped cactimay occur under a loose canopy layer or intermingledwith shrubs in denser communities. They are often seen atthe edges of cliffs and exposed rock surfaces together withglobular cacti.

The dry limestone shrubwood, also known as thornscrub, is considered to be the most common andcharacteristic lowland formation of the West Indies.Sometimes restricted to the coast, it may also extend far

Opuntia macracantha, Guanthnamo, Cuba.Vulnerable.

101

inland in many of the Caribbean islands. Spectacularcoastal benches with dry shrubwoods occur at the easternend of Cuba arround Punta Maisi, and a series of upliftedcoastal benches are found on the southern coast ofHispaniola extending inland to considerable altitude. InMaisi it is common to see Melocactus acunai in thinlycoppiced rock flats between microphyllous shrubs stronglymodified into grotesque windswept aberrant forms.Extensive unbroken stands of this vegetation type arefound in the southern terraced Cuban coast betweenMaisi and Cabo Cruz, with the endemic cactus speciesPilosocereus brooksianus, Leptocereus maxonii, L. sylvestris,and Opuntia macracantha, and in north-eastern Cuba,where Leptocereus santamarinae and other species occur.Smaller stands of this vegetation type are found in thenorthern coastal zone of west Cuba, and in south-centralCuba. In Hispaniola there is also an extensive stand inareas below sea level in the Enriquillo Valley Cul-de-sacarea, giving way to succulent sea coast vegetation withMelocactus lemairei, Leptocereus weingartianus, andMammillaria prolifera, at the shores of the lake. Smallerfloras of endemic cacti, other succulents, and spiny shrubsexist along the leeward coast of Jamaica and Puerto Rico,and also in the Bahamas, the Virgin Islands, and LesserAntilles.

2) Semi-desert cactus scrub - This type of openvegetation is characterised by small trees and shrubs withmany succulents, mainly cacti, which are co-dominant oreven dominant in both shrub and canopy layers. The mostconspicuous elements of this vegetation type are the largecolumnar and treeshaped cacti, which are represented bylocal vicariant endemics in each island of the GreaterAntilles and the southern Bahamas. The semi-desertvegetation in the coastal and subcoastal belts in theCaribbean islands is conditioned by an arid climate withnine to eleven dry months and 30-60 cm of annualprecipitation.

In eastern Cuba the open cactus scrubs form anunbroken stretch along the coast from Guantanamo Bayto Imias. From Imias to Maisi smaller fragments occur,especially on the sand deposits of the coastal areas.Several geographically separated associations occurdepending on whether the soil is rocky or sandy and onthe duration of dry periods. On sandy soils Stenocereushystrix, Opuntia dillenii, 0. hystrix, 0. militaris, andPereskia zinniiflora are dominant, while on rocky habitatsDendrocereus nudiflorus, Pilosocereus brooksianus, Harrisiataylori, Melocactus acunae, and Agave albescens are morefrequent. In the Dominican Republic and Haiti, in theEnriquillo Valley and Cul-de-sac area, there is animpressive cactus scrub with the arboreal Leptocereuspaniculatus and Opuntia moniliformis. It also occurs onalternating salt rock and gypsum outcrops on the slopes ofthe abutting mountain ranges, and in the Azua andBayahibe areas. In Haiti, in the north-western peninsula,there is another stand with the local endemics Opuntia

Opuntia moniliformk in coastal scrubwoods, north-west Haiti.

falcata, 0. ekmanii, and 0. acaulis. A spot of thisvegetation type occurs in Mona Island, Puerto Rico.Dominant cactus species in Mona are Pilosocereus royenii,Stenocereus hystrti, and Harrisia portoricensis.

3) Dry serpentine shrubwoods - The vegetation isdominated by a dense, 2-4 m high, closed shrub layer,small emergent palms, dwarf palms and 4-6 m highmicrophyllous evergreen trees. These dense stands usuallyalternate with small grassy clearings which are oftentransformed into dwarfgrass savannas by humaninterference and grazing. This vegetation type, which iswell developed in Cuba along the central portion of themain island on ophiolitic rock outcrops, is rare or non-existant elsewhere in the Caribbean Islands. Unlike thedry shrubwoods on limestone in Cuba, the serpentinecommunities are physiognomically quite uniform despitethe great differences in climate and floristic compositionamong the outcrops. They are also relatively devoid ofcacti, with the exception of five rare Melocactus speciesand a unique species of endemic Escobaria, all of whichare highly endangered.

Melocactus actinacan thus, 111. matanzanus, and M .guitartii occur in the scrubs of western-central Cuba, whileM. holguinensis, M. radoczii, and Escobaria cubensis areonly found in eastern Cuba.

III. Montane formations1) Tropical karstic forests - These are limestone-basedforests composed of primarily deciduous species withseasonal flowering. They are only found on the ‘mogotes’or haystack mountains of Cuba and Puerto Rico, theCockpit Country and the John Crow Mountains inJamaica, and on the Samana Peninsula of Hispaniola. Ofthese areas, only the Cuban mogotes have proven tosupport a noteworthy succulent flora. The mogote karsticforests have two evolutionary centres in Cuba withdifferent floras, the oldest and richest (40 per centendemic) in the western part of the island, and theyoungest in central and eastern Cuba.

102

The western mogote forests occur on bare rocks ofdeeply eroded mountains and solitary cliffs consistingmainly of hard crystalline limestone. There is a single 4-9m open canopy layer underneath which many smallerplants thrive thanks to the favourable light conditions.Bombacopsis cubensis, a Cuban endemic tree with barrel-like trunk capable of water-storage, is present along withvarious species of Leptocereus with very restricteddistribution patterns, including L. assurgens, L. ekmanii,L. Zeonii, and L. prostratus. Other cactus genera occurringin this habitat are: Harrisia, Selenicereus, Opuntia, andRhipsalis. Agave tubuZata is only found on the cliffs of thewestern mogotes. In the central and eastern karstic foreststhere is only one local species of Leptocereus, L. carinatus.

The most important areas where cacti and succulentsoccur in the West Indies are listed below in the sectionPriority Sites for Conservation.

ThreatsIsland floras are usually composed of ecologicallyrestricted populations not capable of adapting to majorenvironmental changes. The ability of the islandcommunities to recuperate from external pressures is, ingeneral, reduced. This characteristic makes themparticularly vulnerable to degradation. There is a geneticreason for the vulnerability of island communities: thereduced gene pool from which the populations thatcolonised the new biotopes are selected. This gene pooldoes not have the opportunity to improve over a longperiod of time.

The ‘island’ effect in the West Indian Archipelago isincreased due to the fact that most of the larger islandfloras consist of groups of ancient, isolated floras (Borhidi1991). As a consequence of isolation, the ecologicaltolerance and the genetic flexibility of populationsdecrease so that the competitiveness of species becomeslow. Therefore, they cannot react satisfactorily to newenvironmental onslaughts, cannot take advantage of

A massive Dendrocereus nudiflorus tree more than300 years old (diameter of trunk at base: 1.30 m),GuantAnamo, Cuba. Vulnerable.

Bat-pollinated flower of Dendrocereus nudiflorus.

succession, and cannot resist, or force back, newcompetitors. The vulnerability of the succulent endemicflora of the West Indies, in particular, is very pronouncedbecause many taxa have adapted to the extremeecological conditions of oligotrophic or bare substrates.Thus, the level of metabolism becomes low in theseplants, and their competitiveness and degree of sociabilitydiminish even more. On the other hand, it is quite likelythat since Pleistocene times the formerly widespreadxerophilous elements of the larger Caribbean Islandsretreated to relict habitats: dry coastal or subcoastalareas, serpentines, and the slopekarsts. This was probably due to cha

and cliffs of conicalnges to a more humid

climate.The relict character of the succulent native flora of the

West Indies - another condition that increasesvulnerability - is indicated by the presence of relativelyprimitive, isolated taxa (e.g. the dioecious Antillean groupof Pereskia, Dendrocereus); the abundance of disjunctgeographical distribution types, for example Dendrocereusnudiflorus, Bombacopsis cubensis, Furcraea hexupetula,Opuntia nashii, and 0. millspaughii; and the large numberof local endemics represented by small populations, suchas Opuntia hystrix, 0. militaris, 0. sanguinea, 0.borinquensis, and Leptocereus spp. Consequently, theWest Indian succulent flora is one of the mostendangered plant communities in the world.

Clearing for agriculture - The human population of theCaribbean Islands in 1991 was estimated to be 35 million(FAO 1990), projected to rise to about 40 million by theyear 2000, and to nearly 60 million by 2025. In general,

z!although all-island totals are increasing, the incremental

=cd rates and proportions of populations economically active7 in agriculture have fallen over the past decade. Thesezg trends reflect increase in mechanisation and efficiency on5 farms and in importation of foods replacing locala produce, combined with migration to towns and coastal

tourist resorts. Higher growth-rates are associated withlarger rural populations and relatively smaller movementsaway from agricultural pursuits, as in Jamaica, Haiti, and

103

the Dominican Republic. The total number of peoplecomprising agricultural communities actually rose inJamaica and Haiti (by 7.5 per cent) during the decade,this trend matching several of the poorer CentralAmerican republics where, however, numbers of thoseactually economically active on the land also fell.

Most agricultural development has been carried outon the seasonal evergreen forest or seasonal rain forestareas, and on lowlands formerly occupied by semi-deciduous forest, which are not succulent-richcommunities. Nevertheless, important coastal shrubwoodareas in western Cuba with Dendrocereus nudiflorus andPilosocereus robinii have been cleared for sisal (Agavesisalana and A. furcroydes) plantations. Traditionalcassava, maize, and other small crop cultivation has alsobeen attempted locally, with limited success, in manyWest Indian scrub areas.

Grazing - The impact of drought-resistant cattle in theseasonally dry vegetation of the West Indies is wellknown. In the eastern lowlands of Cuba many stands ofsemi-deciduous xerophytic forests with Pereskia zinniifloraand Harrisia sp. were cleared for grazing. Goats areusually grazed in the succulent-rich shrublands of south-eastern Cuba, north-western Haiti, and southernDominican Republic. Opuntia caribaea which isconsidered a pest in Hispaniola, is one of the few nativesucculents that seems to benefit in overgrazed areas.

Burning - The natural vegetation of the West Indianislands has not evolved the fire-tolerant life forms that arefound throughout much of tropical Africa or continentalAmerica, so fire is comparatively more destructive in theCaribbean. Fire is commonly used to clear land foragriculture and settlements, to ‘clean’ undergrowth inforests and to encourage new growth in savannas andbushland in the dry season for pasturage. Some of thearboreal Opuntia (subgenus Consolea), with thick-barkedtrunks, are more fire-tolerant than smaller cacti and canescape damage from fire to some degree, and so canEscobaria cubensis which in the dry season is at groundlevel or even below. But in general, burning is a majorthreat to the succulent plant flora. Particularly susceptibleto fire damage are the thin-stemmed Harrisia spp. andmost Melocactus spp. Harrisia portoricensis became extinctin the main island of Puerto Rico mainly because of fire;now the species is confined to Mona and Monito Islands.Opuntia borinquensis, which has not been seen in the wildfor many years, is probably gone forever due to severeburning of the Cabo Rojo area in south-western PuertoRico, some years ago. This place was considered animportant regeneration area for Melocactus intortus inPuerto Rico; today very few individuals of this species canbe seen there.

Urbanisation and tourism - There are 16 urban centres inthe Caribbean with over 100,000 inhabitants. These are

Cuba: Havana, Camaguey, Santiago de Cuba,Guantanamo; Dominican Republic: Santo Domingo,Santiago de 10s Caballeros; Haiti: Port-au-Prince; PuertoRico: San Juan; Jamaica: Kingston, Montego Bay;Bahamas: Gran Bahama, Nassau; Guadeloupe: Pointe-a-Pitre; Martinique: Fort-de-France; Barbados:Bridgetown; Trinidad: Port-of-Spain. Most of thesecentres are located in coastal areas.

Many restricted endemics have suffered the locationof an urban centre within their sites of occurrence.Escobaria cubensis is facing the growth of the city ofHolguin in eastern Cuba, and is dangerously retreating.Borrichia cubana and Leptocereus wrightii, formeroccupants of the rocky coastal areas of Havana harborand its environs, are nearly extinct now. Agave legrellia~taand Pilosocereus robinii are significantly reduced inoccurrence by the urbanisation of north Havana province.But these are not the only victims of the relativelypopulous capital of Cuba; Cnidoscolus quirzquelobatus andC. pagans, both endemic to the Havana city area, seem tohave disappeared forever. They have not been seen in thewild for more than half a century.

Wildlife supported by succulents: endemic arborealrodent (Capromys pilorides) on columnar cactusStenocereus hystrix, Guanthamo, Cuba.

104

The growth of tourism in many of the islands over thepast 50 years has resulted in hotel development alongcoastlines that have attractive sandy beaches. This hasoften meant a complete change to the landscape locally,involving the removal of natural vegetation and theplanting of ornamental trees, shrubs, and grass for lawnsand golf courses. Varadero Beach in Cuba and LaRomana touristic complex in the Dominican Republic areclear examples of this landscaping policy. The giantDendrocereus nudiflorus in Varadero’s famous beach andresort area has been progressively replaced by RoyalPoncianas (Delonix regia) and other foreign floweringtrees. The dioecious Pereskia quisqueyana, endemic to theBayahibe beach and coastal lowlands in south-easternDominican Republic, is practically extinct by now; veryfew male individuals were left on the strand and as yetnobody has ever seen a female plant.

Mosquito control and marina developments haveeliminated mangroves and littoral thickets in many places.New roads have often been constructed to give access tocoastal areas, and to connect cays with the mainland.

In some Caribbean islands the movement of peoplefrom rural areas to towns and resorts with the lure ofemployment opportunities in servicing the new tourismhas coincided with the decline of export-based plantationagriculture. Increased demand for fresh fruit andvegetables has often resulted in unacceptable levels ofcultivation on unsuitable land.

Mining and quarrying - Threats to the landscape in theWest Indies arise mostly from mining activities. Whatevertype of mining is carried out, vegetation is cleared andthere is always some surface disturbance either fromstripping operations or dumping of tailings. Theexploitation of bauxite in south-western DominicanRepublic has had a significant impact on the dryshrubwoods of the Pedernales area. Quarrying forlimestone in the haystack karstic mountains of north-eastern Puerto Rico and western Cuba has producedsignificant changes in local landscapes and strain onvarious species, notably Leptocereus leonii and L .scopulophilus.

Riversides and beaches are often exploiteddestructively for building sand and gravel and theseremovals may have secondary effects in the form oferosion, flooding, pollution, and loss of visual aesthetic.

Collecting for horticulture - Some of the cacti andsucculents of the Caribbean region are of considerablehorticultural interest and some have been exploited to thedetriment of wild populations. Undoubtedly, the mostdemanded genus is Melocactus. M. intortus has beenheavily collected in Puerto Rico, for example (Martin andFarmer 1975) where hundreds of plants were formerlyremoved from the wild. It has also been exported inquantity from St. Eustatius and Grenada (Howard 1977).Fortunately, M. intortus is the most widespread andabundant species of this genus in the West Indies.

Melocactus lemairei of Hispaniola, M. harlowii ofeastern Cuba, and especially the ophiolitic dwellers M.matanzanus, M. actinacanthus, and M. guitartii wereheavily collected in the 1970s according to local fashionsin their respective countries of origin. Many peoplewanted rock gardens at that time. In Cuba the demand forM. actinacanthus led to the wholesale removal of nearlyall the wild population; only a few dozen individuals canbe seen nowadays in their original location, in very steep,nearly inaccessible cliffs.

The main Caribbean country involved in the export ofcacti and other succulents for horticulture is theDominican Republic. The bulk of the plants exported arenon-indigenous, commonly cultivated Mexican cacti, Aloeand Euphorbia spp. Small quantities of indigenous cactihave, however, been exported including Opun tia an tillana,Harrisia divaricata, and Melocactus lemairei (Oldfield1991). One large commercial outlet in the country exportsmainly to the US.

Introduced species - The introduction of the extremelyaggressive shrub Dichrostachys cinerea from the savannasof Africa into Cuba and Marie Galante has been anecological disaster. The plant regenerates vigorously fromthe smallest root fragments. In south-central Cuba thisinvader has had a notable detrimental impact onLeptocereus arboreus and the xerophytic indigenousvegetation with which this species is associated.

On the small island of Guana in British Virgin Islandsinvasive succulent plants, originally introduced forhorticultural purposes, include Sansevieria trif&-ciata,Kalanchoe spp., and AZoe Vera. These pose a significantthreat to the native flora and are now being controlled(Krauss 1991).

Local use - The fruits of various species of Opuntia,Harrisia, P i l o s o c e r e u s , Melocactrts, HyIocere~rs,Selenicereus, Pereskia aculea ta, and Stenocereus hystrix areused sporadically as sources of food in most West Indianislands, though none of them is commercialised. In north-eastern Cuba a traditional red wine is made out of theripe fruits of Opuntia dillenii. The sticky mucilage fromthe stems of this species, mixed with slaked lime, is still inuse in Cuba to make a coarse paint. Stenocereus hystriu isalso used in Hispaniola, Cuba, and Jamaica as livefencing.

Agave sobolifera and other species are used for craftmaterials in Jamaica and other Caribbean islands. Thesaponin-rich juice extracted from the leaves of Agavealbescens and A. underwoodii serves locally as a laundrydetergent in times of needs. It is not yet well known towhat extent these uses have a detrimental impact on wildpopulations.

Natural environmental factors - Hurricanes, volcaniceruptions, severe droughts, floods, and landslides areintrinsically associated with the West Indianenvironments, and may have profound effects on the

105

Leptocereuspaniculatus, DominicanRepu blk.

floras. On April 13, 1979, for example, the rareSelenicereus innesii seems to have been completely wipedout of the wild, when the area it inhabited in St. Vincentwas devastated by the violent eruption of La Soufriere.

Conservation status

In general, it remains difficult to estimate population sizeand area of dispersal for rare plant species of theCaribbean Islands. Gaps in the flora coverage and thelimited number of people with specialised knowledgehave restricted the availability of data on the conservationstatus of West Indian plants. Very preliminary andincomplete assessments of the cacti and other succulentshave been made. Taxonomic uncertainties add to thedifficulties of applying conservation categories. WCMChas records of over 100 Caribbean cacti in its PlantsDatabase. Of these, about 40 are threatened on a worldscale.

It has not been until now that a more realisticevaluation of the conservation status of the West Indiancacti has been attempted, and it provides the basis of apreparation of a systematic treatment of the Cactaceaefor the New York Botanical Garden’s Flora of theGreater Antilles project. Based primarily on my fieldknowledge of the West Indian islands I have prepared thelist of 243 succulent plant species in 16 families endemicto the West Indies (Table 3 of Annex 14). As noted aboveAruba, Bonaire, Curacao, Trinidad, Tobago, Margarita,and other small islands adjacent to Venezuela wereexcluded from the survey due to the fact that theseterritories are phytogeographically part of northern SouthAmerica, and their flora predominantly continental. Anappreciation of the current conservation status of eachtaxon is given by classifying them within four categories.

For some Haitian rare succulent species with very

limited distribution data and doubtful assessmentinformation on which to base current conservation status,it is necesary to carry out field surveys for more accuratefield-based assessments of their conservation condition.


The West Indian endemic succulent flora occursthroughout the many islands of this dispersed archipelago.Nearly every island, or group of islands, has its own localtaxa. Rather than a few large protected areas, thecoverage of such a disperse flora, in terms of conservationneeds, requires a number of small-sized sites:

Cuba:1) Coast and lowlands from Guantcinamo Bay to Punta

Maisi: an important part of the south-eastern dryshrubwoods and semi-desert cactus communities withAgave spp., Plumeria spp., Cissus spp., and 18 speciesof Cactaceae distributed in 11 genera. This area canbe identified as particularly outstanding for succulentconservation in the Caribbean.

$=IZE<4

Coastal limestone habitat with Stenocereus hystrix,Guanthnamo Bay, Cuba.

106

4

3)

4)

5)

6)

7)

8)

9)

Coast and lowlands of Baconao, sections El Indio-ElMorrillo (Reserva de Santiago): another part of thesouth-eastern dry scrubwoods and semi-desert cactusscrubs including the type locality of the rareLeptocereus maxonii.Coastal section between Pilon and Cabo Cruz: also insouth-eastern Cuba. With some of the succulentspecies of the above mentioned areas, it also includesthe entire site of occurrence of Leptocereus sylvestris.Coastal terrace between Gibara and Puerto Padre bay:an important area of the north-eastern xerophyticplant communities. An undescribed subspecies ofOpuntia nashii grows in this site, together with thegiant Dendrocereus nudiflorus.Coca, Paredon Grande, Roman0 and Sabinal cays:important Leptocereus santamarinae populations occurin some of the largest islets of the CamagueyArchipelago in northern Cuba. The cays are alsoinhabited by Opuntia millspaughii, Pilosocereusmillspaughii (at the southern end of its distribution)and the rare Selenicereus brevispinus, a very restrictedlocal endemic.Santa Cruz de1 Sur lowlands: the savannas north toSanta Cruz, in southern Cuba (Camaguey province),have the largest populations of Pereskia zinniiflora ofboth sexes, male and female. This Cuban endemic isthreatened by habitat destruction at its otherlocations.Coast and lowlands from Playa Giron to Cienfuegos Bay,and from Cienfuegos to Trinidad: important for thelocal endemics Agave acicularis, A. grisea, a n dLeptocereus arboreus. The site also has Pilosocereus,Harrisia, Dendrocereus, Pereskia, Selenicereus,Hylocereus, and Opuntia species.Eastern tip of Hicacos Peninsula, from Rincon Frances tothe east end: the most outstanding area of north-western xerophytic communities in Cuba. Importantfor populations of Pilosocereus robinii, Agavelegrelliana, and Omphalea trichotoma. The site also hasDendrocereus.Guanacahabibes Peninsula: part of the western dryshrubwoods with the rare Harrisia taetra.

10) Western haystack mountain complex of Sierra de 10sOrganos: the whole karstic mountain range, in Pinarde1 Rio province, should be an internationalconservation priority. It has mostly primary vegetationwith Bombacopsis cubensis, Omphalea hypoleuca,Cnidoscolus spp., Cissus spp., Agave tubulata, and 11species of cacti, two of which are yet undescribed.

11) Sierra de Anafe: an isolated limestone mountain rangein Havana province, with Leptocereus leonii.

12) Sierra de Somorrostro and neighboring hills: this othersite in Havana province is important for the highlyendangered Leptocereus scopulophilus.

13) Sierra de Najasa: an interesting site in Camagueyprovince with Hildegardia cubensis and Leptocereuscarina tus.

14) Jibacoa limestone-mountain area in Guamuhaya: steepcliffs inhabited with the local endemic Melocactusperezassoi.

15) Dry serpentine shrubwood sites on rocky outcrops of theophiolitic complex: there are four geographicallyunrelated sites each of which has its own endemictaxa, mostly occurring in small, restricted populations.These are:a) Tres Ceibas (Havana prov.) with Mdocactr~s

matanzanus,b) Agabama (Vi l l a C la ra p rov . ) wi th M.

actinacanthus,c) Jatibonico (S. Spiritus prov.) with M. guitartii,d) Holguin (Holguin prov.) with M. holguinensis and

Escobaria cubensis.

Hispaniola:

2)

3)

4)

5)

6)

7)

Lake Enriquillo Valley, encircled by Jimani-LaDescubierta and Neiba-Duverge (Dominican Republic):a very important part of the south-western semi-desertcactus communities dominated by Leptocereuspanic&a tus, Stenocereus hystrix, and Opuntiamoniliformis. Because of its cactus richness anddiversity (9 genera, 14 species) this area is ofparticular interest for succulent plant conservation inthe Caribbean. Isla Cabritos National Park, a smallisland within Lake Enriquillo, gives insufficientprotected coverage to this unique xerophilous flora.Dry shrublands between Bani and Azua (DominicanRepublic): another part of the cactus scrub ofsouthern Hispaniola, with six different species ofOpun tia.Coast and lowlands between Mole St. Nicolas and Port-de-Paix (Haiti, Dept. du Nord’ Ouest): an importantarea of the north-western xerophytic communitieswith the local endemics Opuntia falcata, 0. ekmanii, 0.acaulis, and a probable undescribed species ofLeptocereus. Knowledge of the status in the wild ofthese taxa is essential to determine their preciseconservation needs.Coast and lowlands of Barahona and Pedernalesprovinces (Dominican Republic): scarcely disturbeddry shrubwoods and semi-desert cactus scrubs withDendrocereus undulosus. Other important cactusgenera present are Opuntia, Leptocereus, Harrisia,Mammillaria, P i l osocereus , Stenocereus, a n dMelocactus.Coast and lowlands between La Romana and CaboEngano, including Saona island (Eastern DominicanRepublic): an interesting part of the southern drycommunities with an undescribed taxon of Opuntia.Dry lowlands south of Montecristi: the most importantarea of the north-western dry shrubwoods and semi-desert cactus communities of the DominicanRepublic.Coast and Zowlands of Bayahibe (Dominican Republic):with the rare Pereskia quisqueyana.

107

8) Cerro de San Francisco, Banica (Elias Pina province,Dominican Republic): important for Pereskiamarcanoi. 2)

Puerto Rico: 3)

3)

4)

Coast and lowlands from Bahia de Guayanilla toBoqueron: the most important part of the dryshrubwoods and semi-deserts of the main island, with 4)12 species of Cactaceae, half of which are Opuntia.Mona and Monito Islands: a sanctuary for the Puerto

Islands: dry shrubwoods with Opuntia millspaughii, 0.bahamana, 0. lucayana, and Limonium hahamense.Eleuthera Island rocky plains: important for Agavcbraceana, A. cacozela, and Pilosocereus bahamensis.Great Inagua open sandy flats and rocky coastal coppices:with Agave inaguensis, A. nashii, and Pilosocereusmillspaughii.Long Island dry shrublands: important for Agavcindaga torum and Harrisia brookii.

Rican xerophytic communities, with the rare Harrisia The Lesser Antilles:portoricensis and a yet undescribed taxon of Opuntia.South-eastern Culebra Island: the only site ofoccurrence of Leptocereus grantianus.Tetas de Cayey mountain ridge: an interesting rockyoutcrop with an undescribed Melocactus taxon.

9

2)

3)Jamaica:

1) Hellshire Hills coastal and lowland area: an importantpart, south to Spanish Town, of the southern dryshrubwoods and semi-desert cactus communities withthe endemics Opuntia spinosissima, 0. jamaicensis,Pilosocereus swartzii, and Melocactus caroli-linnaei.Coast and lowlands from Treasure Beach to Little PedroPoint area: another part of the southern dryshrubwoods with Acanthocereus sp., in St. Elizabethparish.

4)

5)

Northern Grand Terre of Guadeloupe: dry shrubwoodswith Agave dussiana, Acanthocereus pentagonus,Opuntia rubescens, and 0. triacantha.Barbuda lowlands: with Agave karatto, Manzmillarianivosa, Melocactus intortus, and 0. rubescens.South-eastern peninsula of St. Kitts: important forAgave van-grolae, Melocactus intortus, and Opuntia spp.La Soufriere volcanic cone, St. Vincent: the only knownsite of occurrence of Selenicereus innesii. This rarespecies could have become extinct after the 1979eruption.Additional sites: Antigua, south-western area;Anegada, eastern lowlands and eastern central area;Virgin Gorda, central coast area; Martinique, south-western area.

Cockpit Country: interesting karstic forests in The Cayman Islands:Trelawny parish, with a rare Mammillaria species on 1) The Bluff area of Cayman Brat: with an undescribedvertical limestone cliffs. variety of Epiphyllum phyllan thus, Opun tia

millspaughii, and new taxa of Harrisia and Pilosocereus.The Bahama Archipelago:1) Coastal areas and rocky plains of the Turks and Caicos

2) East End area of Little Cayman: part of the cactusscrub of the Caymans, with Opuntia, Harrisia, andPilosocereus.

Table 3.17 Caribbean protected areas: number and area covered (in hectares) by IUCNCategory Category definitions are given in Box 2.4.

Country Cat. I Cat. II Cat. Ill Cat. IV Cat. V TOTAL

Antigua & Barbuda 114,128 114,128Bahamas 4/l 21,516 l/l ,813 5/l 23,389Belize l/4,144 6/l 13,846 7/l 17,990Bermuda 1/12,000 1/12,000Caymans l/1,731 l/3,310 215,041Cuba 8132,405 8199,518 21/l 70,487 36/l ,038,996 73/l ,341,406Dominica 116,872 116,872Dominican Republic 1 O/488,069 7/476,090 17/964,159Guadeloupe 1 /I 7,300 l/3,700 2/21,000Haiti 217,500 112,200 319) 700Jamaica 2/37,953 2137,953Martinique I/ 70,150 I/ 70,150Dutch Antilles 217,760 217,760Puerto Rico 14128,548 14128,548St Lucia l/1,494 111,494St Vincent &Grenadines 218,284 218,284Trinidad &Tobago 7115,528 7115,528Turks & Caicos 10186,745 II1259 l/4,497 4/5,301 14197,532BVI 3/673 31673USVI 1114,079 1114,079

108

Existing conservation measures Cabo Corrientes and Baconao reserves (Cuba); lsla

Unfortunately, until very recently the conservation of theCaribbean biodiversity was not recognised as being ofimportance by international conservation agencies. Therehas been remarkably little attention paid to theconservation needs of the cactus and succulent flora ingeneral. Undoubtedly, the historical preoccupation withtree forest, and especially our present-day concern fortropical rain forests, have diverted attention from othervegetations of equally deserving botanical merit in theCaribbean (Adams 1997). Plant formations in need ofprotection include special floras of serpentine soils inCuba and Jamaica, and siliceous sand savannas whichhave a restricted range outside Cuba. Recently attentionhas been concentrated on the establishment of marineprotected areas. These may incidentally protect thesucculent flora of coastal areas (e.g. Buck Island, in St.Croix, US Virgin Islands) but do not necessarily protectthe habitats of endemic and/or threatened succulentspecies.

Sometimes areas of cactus scrub vegetation areprotected within various sites set aside for conservationthroughout the Caribbean. In the British Virgin Islands,for example, the Fallen Jerusalem and West Dog IslandForestry Parks support this type of vegetation. Somepriority sites for the protection of specificallyrepresentative areas of cactus scrub in the West Indies are

Cabritos National Park (Dominican Republic); GuanicaState Forest and Mona Island reserve (Puerto Rico), andPart National (proposed in Guadeloupe).

Protected areasProtected areas currently cover not more than 6.6 percentof the Caribbean Islands’ land area, and West Indianbiodiversity is not fully represented. Besides, many of theprotected areas which do exist are not adequatelyprotected on the ground. According to an AS0 study(Inventory of Caribbean Marine and Coastal ProtectedAreas, 1988) only 33 percent of the protected areacoverage is under a suitable regime of protection andmanagement; 43 percent is partially protected, while theremaining 24 percent is practically unprotected. A reviewof the protected areas system and conservation legislationin the Caribbean is given in IUCN (1992). A summary ofprotected areas in the West Indies is presented in Table3.17, and specific areas important for succulentconservation are presented in Box 3.6.

Notwithstanding that the most common of the WestIndian lowland formations is the thorn scrub, most of thearea coverage listed in Table 3.17 is of montane andsubmontane rain forests, seasonal evergreen forests, semi-deciduous, and coniferous forests.

Box 3.6 Protected areas important for succulent conservation in selected Caribbeanislands

Cuba

Area (km*) Category

1) Gran Parque Sierra Maestra Integrated Management Area comprising:a) National Park of Desembarco del Granmab) National Park of Turquinoc) Biosphere Reserve of Baconao

2) Escambray Integrated Management Area3) Mil Cumbres Integrated Management Area4) Vinales National Park5) Peninsula de Guanacahabibes Biosphere Reserve6) Cabo Corrientes Natural Reserve7) Subarchipielago Sabana-Camaguey Integrated Management Area8) Punta Frances-Punta Pedrales Natural Park

5,270 VIII258 II175 II846 IX

1,870 VIII166 VIII134 II

1,015 IX16 I

179 V174 II

Hispaniola1) lsla Cabritos National Park2) Del Este National Park including Saona island3) Monte Cristi National Park4) Sierra de Bahoruco National Park5) Jaragua National Park including Beata island and the submerged platform

between the island and the mainland)

24 II420 II

1,309 II800 II

1,374 II

Puerto Rico1) Guanica State Forest2) Mona Island Natural Reserve3) Culebra National Wildlife Refuge

JamaicaI) Hellshire Hills (proposed) reserve2) Cockpit Country

55 IV6 IV

109

Conservation agencies and botanical institutionsCuba

1)

2)

3)

4)

5

6)

7)

Viceministerio Forestal, Ministerio de Agricultura(MINAGRI): This government institution hasresponsibility for forest management and protectedareas. MINAGRI is also the CITES ManagementAuthority.Comision para la Protection de la Flora, la Fauna y elus0 rational de 10s R e c u r s o s N a t u r a l e s(COMARNA): This government entity, which worksclosely with MINAGRI, investigates the developmentof management plans for protected areas.Instituto de Ecologia y Sistematica (IES): Researchinstitute under the Ministerio de las Ciencias y laTecnologia (former Academia de Ciencias), with acore of botanical and zoological staff supported by astrong reference library and the largest herbarium ofthe country. It includes the Centro National deBiodiversidad which is developing a database on WestIndian biodiversity.Jardin Botanic0 National (JBN): Large garden underthe Universidad de La Habana (Facultad de Biologia);it maintains important collections of Cuban plants.There is also a core of botanical staff and an importantlibrary.Museo National de Historia Natural (MNHN): Alsounder the Ministerio de las Ciencias y la Tecnologia,with a group of well trained and experienced curators,one of which is a Scientific Advisor of RARE Centrefor Tropical Conservation.Centro Oriental de Biodiversidad y Ecosistemas(COBE): The same as IES in the eastern provinces.Instituto Superior Pedagogic0 (ISP): Important groupof institutes (one in each province) under theMinisterio de Education, with a Department ofBotany and staff specialised in local (provincial)floras.Sociedad pro-Naturaleza (SPN): Recently establishedNGO for the protection of the environment throughlocal initiatives.

l * - 110Dommlcan KepuwcDirection National de Parques (DNP): Thisgovernment body, under the Secretaria de Estado deAgricultura (SEA), Subsecretaria de RecursosNaturales, is responsible for forest management andprotected areas. DNP is also the CITES ManagementAuthority.Jardin Botanic0 National “Rafael M. MOSCOSO” (JBN-RMM): Maintains collections of Hispaniolan plants.At the herbarium there is a small but experiencednumber of curators.Departamento de Biologia, Universidad Autonoma deSanto Domingo (UASD): with a Biological staff, andherbarium.Parque Jaragua (PJ): An NGO for the study andprotection of wildlife in Parque National Jaragua. It

also investigates the development of managementplans for this protected area.

5) Natura (N): Another NGO which serves to checkenvironmental health. Provides control activitiesdeveloped through local initiatives.

Puerto Rico

1)

2)

3)

4)

5)

6

Departamento de Recursos Naturales (DRN): Thisgovernment Department has responsibility for forestmanagement and protected areas. DRN is also theCITES Management Authority.Jardin Botanic0 de la Universidad de Puerto Rico,Recinto de Rio Piedras (JB-UPR): Maintains livingcollections of Puerto Rican plants and a herbarium.Centro de Information Ambiental de1 Caribe (CIAC):Recently established centre under the UniversidadMetropolitana, to develop a database on Caribbeanenvironment concerns.Centro de Education Ambiental (CEA): Alsoestablished under the Universidad Metropolitana, topopularise environmental information witheducational purposes.Fundacion Puertoriquena para la Conservation(FPC): An NGO for the conservation of the island’sflora and fauna.Fideicomiso para la Conservation (FC): AnotherNGO for the rescue and protection of Puerto Rico’snatural and cultural heritage.

.Jamaica

1)

2)

Natural Resources Conservation Department(NRCD): Government Department responsible forforest management and protected areas. It is also theCITES Management Authority.Institute of Jamaica (IJ): Renowned Jamaicaninstitution with the most valuable herbariumcollections of the Caribbean.Department of Botany, University of West Indies(UWI): With an experienced staff, and herbarium.Entity: Encourages economic progress coupled withnatural resource protection through education andhosting a tree bank.Natural History Society of Jamaica (NHS):Disseminates information and provides education onconservation and preservation of the environment.Jamaican Society of Scientists and Technologies:Supports scientists and resource development.

National and regional legislationSome Caribbean islands have national legislationprotecting the environment (e.g. Law no. 33 of 12 July1981, in Cuba). Although rare and threatened plantspecies are considered in some of these legislations, ingeneral legal protection for succulents has not been fullyelaborated. The SPAW Pro toco l , ou t l ined inInternational legislation in Chapter 2, if effectivelyimplemented may provide the impetus to update ordevelop new legislation.

110

Ex situ conservation and reintroductionFor a number of highly threatened succulent species thereis no other alternative, at this moment, than to conserveand propagate them ex situ. Studies need to be carried outto find similar habitats for replanting, following the IUCNGuidelines for Re-ln@oductions (IUCN 1995). Examples ofsmall-scale reintroduction projects for cacti have takenplace in Cuba, Puerto Rico, and Guana, British VirginIslands.

South AmericaNigel Taylor, Roberto Kiesling, and Robert Kraus

The South American Continent (i.e. Panama southwards,including the Caribbean islands of the Dutch Antilles,Trinidad and Tobago, and those belonging to Venezuela,and the archipelagos of the Galapagos and Fernando deNoronha) has a complete spectrum of environments andan extremely large and varied flora numbering in theregion of 90,000 species of higher plants. Of these,succulents probably represent about 1.5 per cent (i.e.< 1400 “pp.), the great majority being endemic and foundin regions with less than 1000 mm of annual precipitation(cactus epiphytes excepted). The principal succulent plantfamilies represented (cf. Eggli 1994), in order ofimportance, are Cactaceae (c. 850 spp. in many genera),Portulacaceae (c. 200-250 often weedy spp. ofAnacampseros, Grahamia, Portulaca, Cistanthe,Montiopsis, Silvaea, Talinum, Xenia), Bromeliaceae (c. 170spp. of Dyckia, Cottendorjia, Encholirium, Deuterocohnia,Abromitiella, etc.), Crassulaceae (c. 30 spp. of Echeveria,V’iZZadia, Sedum etc.), Piperaceae (c. 20 spp. of succulentPeperomia), and less than 10 species in each of Agavaceae(Agave, Furcraea), Aizoaceae (mostly introduced weeds),Asclepiadaceae (Marsdenia sessilifolia, M. megalantha,Brazil), Bombacaceae (Cavanillesia, Ceiba), Caricaceae(Jacaratia corumbensis, Carica chilensis), Compositae(Duseniella pa tagonica, Argentina), Cucurbitaceae(Apodanthera, some Brazilian spp. only), Dioscoreaceae(Dioscorea basiclavicaulis, Brazil), Euphorbiaceae(Euphorbia spp., Jatropha spp.), Nolanaceae (NolanasPP*), O x a l i d a c e a e (Oxalis spp . , Ch i l e ) , andPhytolaccaceae (Phytolacca dioica). There are also variousspecies of Basellaceae (Boussingaultia “pp.), Begoniaceae(e.g. Begonia venosa), Bromeliaceae (e.g. Puya, Tillandsia,etc.), Commelinaceae (CaZZisia, Tripogandra), Compositae(Senecio), Ericaceae (Sphyrospermum spp.) ,Euphorbiaceae (Euphorbia lutzenbergeriana and allies,Manihot spp.), Gesneriaceae (Nematanthus gregarius,Sinningia leucotricha), Melastomataceae, and hydrophilicCrassulaceae (Tillaea), etc., which are borderlinesucculents, caudiciforms, or waterplants not counted here.

The taxonomic status of species in certain genera ofCactaceae, Portulacaceae, and Bromeliaceae remainsdoubtful, many being poorly or rather narrowly defined.

Succulent plants in the floras of Peru and Bolivia areinadequately understood, even though a modern checklistis available for Peru. There is no overall Flora coveringSouth America, and some of its larger countries, such asBrazil, have no modern Flora of any kind and even lackcompleted Floras at state level. However, floristicknowledge in Argentina, Ecuador, Venezuela, andParaguay is better, there being either comprehensive stateFloras already published or regional/national accounts inpreparation, or with parts including succulents publishedor in finalised manuscript. The largest family involved, theCactaceae, is relatively well-understood in Venezuela,Chile, Argentina, Paraguay, and eastern Brazil, butpresents considerable problems elsewhere.

Takhtajan (1986) divides South America into sixfloristic regions, which are employed in slightly modifiedform below: (1) the Caribbean Region, comprising itssouthernmost islands (CuraGao to Trinidad, etc.), Panamaand the coastal areas of northern Venezuela, Colombia,and north-western Ecuador; (2) the Andean Regionranging from the vicinity of Caracas (Venezuela) andSanta Marta (Colombia) southwards to north-westernArgentina and northern Chile (including east and westmargins of the Atacama Desert), taking in the Pacificcoast from southern Ecuador southwards; (3) the AmazonRegion including the lowlands of the Rio Orinocodrainage; (4) the Region of the Guyana Highlands ofVenezuela, northernmost Brazil, and the Guianas; (5) theBrazilian Region of extra-amazonian Brazil, Paraguay,eastern Bolivia, and northern Argentina (Chacovegetation); and, lastly, (6) the Chile-Patagonian Regionincluding Uruguay and the remaining parts of Argentinaand Chile.

A few widespread and common species of Cactaceaeare probably not at risk and need not be mentioned belowunder individual regions. These include Pereskia aculeata,Rhipsalis baccifera, Disocactus amazonicus, Epiphyllrrmphyllanthus, Pseudorhipsalis ramulosa, and Selenicercussetaceus. The very humid Regions (3) and (4) are oflimited importance for succulent plants (includingepiphytes), these being represented by a few species ofCactaceae (Melocactus, Cereus, Hylocereus, Pilosocereus,Selenicereus), Bromeliaceae (especially >20 spp., GuianaHighlands), and Portulaca, which are either believed tohave extensive distributions or whose habitats are notparticularly threatened by anthropogenic change (e.g.tepuis, inselbergs, riverine rocks, flooded forest). They arcnot considered further here.

The Caribbean Region

The South American part of this region has two basickinds of environment in which succulents are found:tropical rain forest and dry seasonal forest including openscrub. The former is the habitat of a few widespreadcactus epiphytes, that are probably not threatened asspecies. There is also the remarkable woody, terrestrialcactus, Pereskia bleo, from Panama and Colombia

(Leuenberger 1986) which again is not threatened(Leuenberger, in Zitt 1993). In Panama its range appearsto include various protected forest areas (WCMC 1992).Table 3.18 lists succulents under threat in Panama. Thedry seasonal forest/scrub, which includes the extensive,Falcon-Lara Depression of Venezuela, south Caribbeanislands and the Galapagos Archipelago, is home to about30 mainly treelike or shrubby cactus species, three or fourAgavaceae, and one or more Echeveuia spp., the majoritybeing widespread and common. However, on themainland endemics of more restricted distribution arerepresented by Agave cocui, Echeveria sp. nov. (Larastate), Cereus fricii (C. russellianus), C. mortensenii, C.horrispinus, and Pseudoacanthocereus (Acanthocereus)sicariguensis, all from northern Venezuela, the latter tworanging into Colombia, where a local endemic,Armatocereus humilis (Rio Dagua valley, westernColombia), is found. One or more of these could becomethreatened if widespread conversion of its habitat shouldtake place, and it is unclear whether any occur within theprotected areas listed for this region (cf. WCMC 1992).

Table 3.18 Succulent species consideredto be under threat in Panama(Source: Villa-Lobos, in press)

Species Status

Acanthocereus tetragonus VAgave angus tifolia RHylocereus costaricensis VHylocereus monacanthus VHylocereus polyrhizus VHylocereus s tenopterus VPereikia aculea ta VPereskia bleo VPereskia guamacho VWeberocereus panamensis VWerckleocereus tonduzii V

The islands of the southern Caribbean are home totwo species of Melocactus with restricted distributions: M.macracanthos (Dutch Antilles, probably not at risk) andM. broadwayi (Tobago, Grenada, and St Vincent, statusunknown).

The Galapagos islands, which have protected status asan Ecuadorian National Park, are home to Talinumgalapagosurn and about seven rare, endemic cactusspecies, comprising two endemic monotypic genera(Brachycereus, Jasminocereus) and the remarkable, gianttree opuntias. Recent fires on Isabella Island have beencause for concern and pressure from tourism is increasing,though to what extent these are affecting xeric habitats isnot clear. Fortunately, attention from conservationists iscurrently being focused on the archipelago (Jervis 1994).

The Andean Region

This is the most important area for succulent plant generaand species in South America, most of the species andmany of the genera being endemic. Habitats including

succulents are very diverse, ranging from coastal fogdesert, where rainfall is almost non-existent, to humidforest, seasonal dry forest and dry alpine vegetation, suchas the paramos of the northern Andes and the Puna ofArgentina, where some succulents occur at elevations inexcess of 4000 m. Unfortunately there are insufficientdata concerning some of the areas of greatest diversity inthis region (especially Peru), but it seems probable thattaxa occurring in regions above 2500 m are generally inless danger than those from lower elevations, wherehuman influence is strongest. However, in parts of thecentral Andes at high elevations continually expandingprimitive agriculture and over-grazing are substantiallymodifying some ecosystems and almost certainly leadingto the endangerment of endemic species.

The Andean Region and those which follow aretreated mainly on a country-by-country basis:

Columbia and VenezuelaThe part of the northern Andes which falls within theterritories of Colombia and Venezuela is not very rich insucculent plant taxa, but there are 2 endemic Melocactusspp. (Taylor 1991b), about 7 endemic Echeveria (c. 4 spp.are awaiting formal description, Taylor and Eggli, ined.),and at least 2 Agave spp. Of these endemic taxa, only 2(Echeveria recurvata s. l., E. venezuelensis) are known to bewidespread, but most of the remainder, although theymay be rare and lacking any kind of habitat protection,are not known to be under particular threat in the dry,precipitous environments they mostly inhabit. Anexception is Melocactus schatzlii, which occurs at relativelylow elevations in habitats near to roads and villages in theRio Chama valley, south-west of Merida (Venezuela), andto the south of Bucaramanga (Colombia).

Melocactus schatzlii, Rare endemic of two valleys,one in north-west Venezuela, the other in northernColombia.

112

Table 3.19 Distributional information for succulents at species level, by region of Peru(Source: Brako and Zarucchi 1993)

Region of Peru Altitudinal range (m) Species Per cent total Endemics

Amazonian O-500 21 6.9 9Andean I 500--l 500 85 28.0 54Andean II 1500-3500 180 59.2 133Andean III > 3500 44 14.5 26Coastal o-1 000 49 16.1 35

EcuadorThe relatively small succulent flora of the part of Ecuadorconsidered here includes some 25 species of Cactaceae(Madsen 1989) about 4 poorly understood species ofEcheveria (currently being studied for the Flora ofEcuador project) and the endemic Peperomia graveozens.Of the Cactaceae, 8 species are endemic and of theseArmatocereus brevispinus, Cleistocactus leonensis, andEspostoa frutescens are rare and restricted to the RioCatamayo valley (together with some other, non-endemicsucculents), and Weberocereus rosei, known from only 2natural sites in Chimborazo and Canar Provinces, isregarded as endangered (Hunt 1992) and is not includedin any protected area. Likewise, there appear to be noreserves that would afford any protection to the speciesrestricted to the aforementioned Catamayo valley. Thenon-endemic Melocactus peruvianus has apparentlybecome extinct in south-western Ecuador, but iswidespread and thought to be abundant in neighbouringPeru.

PeruThe part of Peru included here has the largest but leastunderstood succulent plant flora in South America and, inthe Americas, in terms of succulent plant diversity Peru issecond only to Mexico. The following discussion utilisesthe annotated checklist of the flowering plants of Peru(Brako and Zarucchi 1993), which accepts 247 taxa* ofCactaceae (accepted and provisionally accepted speciesmainly according to Hunt 1992), 75 per cent said to beendemic; 26 species of Portulacaceae, 35 per centendemic (especially Cistanthe); 20 species of Crassulaceae(Tillaea and introductions excluded), 50 per cent endemic;and one non-endemic succulent Bromeliaceae,Deuterocohnia longipetala. In addition, of the manyspecies of Peperomia known from Peru, at least 10 aresucculent (3 endemic), and there are various endemicNolana spp., but it is uncertain which of these qualify assucculents. There appear to be no endemic Aizoaceae orAgavaceae.

It is unclear whether any of Peru’s endemic succulentsare included within the few protecte-d areas indicated inWCMC (1992).

* At least 13 taxa of Cactaceae are almost certainly synonyms

and 3 spp. have been recorded in error (see Neoporteria), while a

further 16 represent interspecific or intergeneric hybrids.

At generic level the following are endemic andemphasise the importance of the Andean Region(especially subdiv. I-II): Calymmanthium (l-2 “pp.,Andean I); Lasiocereus (2 spp., genus of uncertaintaxonomic status, Andean I-II); Matucana (22 “pp.,various only provisionally accepted, Amazonian andAndean I-III); MiZa (l-3 spp., Andean I-II and Coastal,probably not threatened); Oroya (l-3 spp., Andean II-III);Rauhocereus (1 sp., Coastal).

As the Table 3.19 indicates, the Amazonian Region ofPeru appears to be the least important in terms ofsucculents, and the majority of the species it holds and itsendemics are found in close proximity to the adjacentAndean I subdivision and do not relate to the vastAmazonian Region in the sense of Takhtajan (set above).There is no information available on the conservationstatus of succulent plants from this region.

Subdivisions I and II of the Andean Region of Peru(500-3500 m altitude) are extremely rich in succulentplant species, and on average two thirds of these areendemic. These subdivisions include dry vegetation withconspicuous, large-shrubby or treelike cacti from thegenera Armatocereus, Browningia, C’~~~istoc.aL.tll,~,Corryocactus, Echinopsis, Espostoa, Haageocererrs, andOpuntia, all of which are credited with numerous species(Hunt 1992; Brako and Zarruchi 1993). The lower-growing, endemic genus Matucana also has the majority.of its species in subdivisions I and II. The number ofrecognisable species in these genera will possibly bereduced by 50 per cent or more once proper taxonomicstudies are accomplished, but the level of endemism is

Browningia candelarib, Quebrada Tinaja, Peru.

113

likely to remain high. As in the case of the previousPeruvian region, very little is known of the conservationstatus of succulents in these subdivisions, but shiftingagriculture and especially overgrazing by livestock, whichare inhibiting regeneration of species such as the treeforming cactus Browningia candelaris (Anderson andKattermann, pers. comm. 1994) may be placing somespecies in danger.

Andean subdivision III, i.e. land above 3500 maltitude, is less rich in succulent species than that at lowerAndean elevations and, as might be expected, the taxa itincludes are markedly different. Highly specialised, alpinespecies from the cactus genera Echinopsis (Lobivia),Matucana, Oreocereus, Oroya, Neowerdermannia, andOpuntia are complemented by various endemic Echeveriaand Cistanthe species, most of whose ranges commence inthe Andean II subdivision. Endemism is slightly lower, atabout 59 per cent, but there may be taxa at risk, at least inthe drier parts of south-eastern Peru, where agriculturalpractices similar to those described for the high Andes ofBolivia are assumed to occur (see below, especially withreference to Neowerdermannia).

In terms of numbers, with 49 species the CoastalRegion of Peru is the third most important of the fivesubdivisions recognised here, but has the second highestratio of endemics (71.4 per cent). However, it is the partwhere most of Peru’s human population is located. TheRegion is divisible into two basic zones, the southern, fog-influenced ‘lomas’ (which represent a continuation of theChilean Coastal Fog Desert, see below), and the northernSechura Desert, which is contiguous with the dry south-western corner of Ecuador. According to recentobservations, habitat alteration, through urban expansionand overgrazing, is a serious problem in areas wherewater availability permits the establishment of humansettlements. The seasonal ‘garua’ or coastal fog supportsannual grasses and other temporary vegetation. Livestockare transported to coastal areas to graze upon theseephemeral plants and nurse plants and juvenile cacti areseverely impacted in some areas (Anderson, pers. comm.1994).

BoliviaThe Andean part of Bolivia can be subdivided into threezones, the high Andes to the west and the lower inter-andean valleys to the east (which descend until the Chacois reached) and, partly between these and expanding tothe north, the very humid Yungas forest.

High Andes. A large part of Bolivia is situated in theregion where the high Andes divide and expand into twomajor mountain chains with the altiplano between (the‘nudo andino’ or Andean knot). The altiplano is a largeplateau surrounded and cut through by volcanic,metamorphic, or in many cases sedimentary mountains,the latter having habitats most seriously affected from theconservation standpoint. The rainfall is greater towards

the northern end of this area and permits dense humansettlements to exist despite the high elevations of theregion (3000-5000 m). The main activity is traditionalsubsistence agriculture, most produce being consumedlocally, and heavy machinery is used only in a few placeswhere there are wider plains of clayey soil. Mostcultivation takes place on steep slopes, which aresometimes terraced. The farmers collect the many stonesfrom the small fields (average minimum of 100 mz), pilingthem up at the field edges, having first removed all thenatural vegetation of scattered shrubs, grasses, otherherbs and cacti. Then the men take the ox-plough to thefield and are followed by the rest of the family who plantcrops such as potatoes, broad beans, corn, andChenopodium quinoa. Each field is used for one year thenleft fallow for 3-7 years, or less if the locality is highlypopulated. The summer rains expose the clay surfacewhich is then eroded by the wind during the dry winterseason. A good example of this agricultural system is thesedimentary plateau at 5000 m elevation near the airportat La Paz. In this region, which presumably continues intothe dry Andes of adjacent Peru, all the vegetation isstrongly affected by human activity. The traditionalagricultural techniques continue to expand into newareas. The perennial vegetation disappears leaving onlyshortlived herbs and shrub seedlings. The cacti, which arevirtually the only succulents of the region, almostdisappear. If one estimates the density of cacti per squaremetre, in a hectare plot about 10,000 plants are destroyed

Lepismium bolivianurn, known only from the Yungasforests of Depto. La Paz, Bolivia.

114

during clearance. If there is an increase of 1000 plots eachyear, then an annual destruction of 10 million cacti can beimagined. The genera with species most affected by thisare Echinopsis (Lobivia, Mediolobivia), Opuntia(Maihueniopsis and the Airampo Group), Corryocactus,Parodia, Rebutia, and Weingartia. Many of their Bolivianspecies are endemics.

Another disturbing development in the high Andes ofBolivia is the planting of Eucalyptus to provide woodwhich is locally in shortage. On the stony slopes where it isnot possible to grow other crops the trees are planted onlytwo metres apart over large areas of land, such as can beseen near Sucre. The shade cast by these plantations killsall the low-growing native vegetation including cacti suchas Echinopsis (Lobivia) spp. and endemic Rebutia(Sulcorebutia) spp. Also of special concern is the status of

Cactaceae at the University of La Paz (by W. Hoffmann etal.). The same university is currently in the early stages ofestablishing a botanic garden, which should help in the exsitu conservation of some of the cacti threatened in thisregion. At present (1994) it does not have facilities for thecultivation of the smaller globular cacti, but has plantedsome of the larger-growing species (Metzing, in Zitt.).

Not all human activities in this region result in anegative influence on the cactus flora. Disturbance orclearance of the natural vegetation sometimes increasesthe abundance of particular species by reducingcompetition, e.g. Opuntia spp., the larger Echinopsis spp.etc., and, in degraded Prosopis woodland, columnar cactiincluding Oreocereus may become more abundant.

Yungas. This narrow band of rain forest extends along theeastern slopes from the central Bolivian Andesnorthwards into Peru and is notable mainly for endemicrepresentatives of the primarily epiphyic Cactaceae tribeRhipsalideae, namely Lepismium crenatum, L.lorentzianum, L. micranthum, L. miyagawae, L.monacanthum, L. paranganiense, L. incachacanum, L.bolivianum, Rhipsalis cuneata, and R. goebeliana. Theconservation status of these cannot be determined atpresent, but most are known from only one or very fewlocalities. Some have been seen recently (e.g. R. cuneata),while others await recollection (e.g. L. miyagawae).

Dry Inter-andean valleys. East of the altiplano ataltitudes of 1000-2500 m there is a dry vegetation whichshares some characteristics with the Chaco. Columnarcactus forests, known as cardonales, are found in the drierparts of this zone, which has a much lower density of

human population than the higher region to the west.Habitat modification is limited to river valleys andincludes primitive cattle ranching (with consequent soilerosion through trampling) and forestry plantations. Inaddition, some of the natural forest on the drier slopes isbeing cleared for charcoal production, all these pressuresbeing of concern to conservationists studying locallyendemic macaw species (M.B. Christiansen, pers. comm.).Treelike and shrubby, endemic cacti commonlyrepresenC. sama

tedipa

in this forest include Cleistocactus parvij?Iorus,tanus, Corryocactus pulquinensis, Harrisia

tetracantha, Neoraimondia herzogiana, Opun tiacochabambensis, Pereskia diaz-romeroana, and P.weberiana. These and other, non-endemic species (e.g. theepiphytic Lepismium ianthothele) may not be underserious threat at present, but the status of more restrictedendemics, such as Samaipat icererrs corroan us,Yungasocereus inquisivensis, Echinopsis (Lobivia)caineana, Gymnocalycium riograndense, Parodiamairanana (P. compressa), and Espostoa (Vatricania)guentheri (Rio Grande drainage system), needs to bemonitored. The first two are monotypic genera, while thelast-named follows river valleys where cultivation is moreprevalent.

NW ArgentinaThe Andes of north-western Argentina comprise thewestern Cordillera de1 Limite and the southern extensionof the Cordillera Real of Bolivia, which, as in the lattercountry, surround the altiplano plateau or Puna. Altitudesrange from 500 to 5000(-6950) m, but succulents areabsent from the higher parts, where cold is too intense,their altitudinal limit reaching 5000 m in Jujuy in thenorth, but down to c. 2500 m in San Juan, and less than1000 m at the southernmost part, to the south ofMendoza, where the climate becomes increasingly wet.

Although succulent species are numerous and manyare endemic, relatively few seem to be of conservationconcern on presen t knowledge. On stony s lopes in the

Blossfeldia liliputana in habitat on cliff in north-westArgentina.

115

Eriosyce aura& after commercial collection inhabitat, northern Chile.

Puna, the attractive, white-woolly, non-endemicOreocereus ~~ollii was formally collected for sale elsewherein the country as a decorative garden plant, but this haslargely stopped with the realisation that most plants soondie. Specimens growing near to roads are still collected byinexperienced tourists, but it remains abundant in lessaccessible areas. Other succulents from this habitat, suchas Anacampseros kurtzii and Parodia maassii, are eithervery common or too insignificant to attract attention fromcollectors.

The dry valleys of the lower Prepuna botanicalprovince have an abundance of succulents from thefamilies Bromeliaceae (Abromitiella, Deuterocohnia,Dyckia), Cactaceae (including the endemic Eriosycesubsect. Pyrrhocactus with 5 spp.), Crassulaceae,Piperaceae (Peperomia) and Portulacaceae, and there arevegetation types dominated by succulents includingtreelike forms (e.g. Echinopsis pasacana). A remarkable,highly specialised, dwarf cactus is Blossfeldia liliputana,but this, like most species, is not significantly threatenedby human influences, and even more local taxa, such asSedum jujuyense (endemic) and Xenia vulcanensis (alsoknown from southern Bolivia), are safe, inhabiting verysteep slopes. Other local endemics include Parodiachrysacanthion, P. penicillata, and P. nivosa, the last-named requiring regular monitoring since a road passesthrough the only locality known.

There is a special conservation problem in themountains of Tucuman and northern Catamarca, wherebroad plains have been cleared for the production of seedpotatoes, the region being free of potato virus. This hascaused the destruction of numerous populations of theendemic Echinopsis (LobivialSoehrensia) bruchii, E.schreiteri (L. stilowiana), and Gymnocalycium baldianum.Fortunately each of these is also found on adjacent stonyslopes that are unsuitable for potato culture, but ourknowledge is incomplete and it cannot be ruled out thatthere are other, more threatened species restricted to theplains favoured for this kind of agriculture. Species thatare abundant on stony ground unsuitable for cultivation

and which can definitely be stated to be out of dangerinclude: Denmoza rhodacantha, Echinopsis (Trichocereus)angelesiae, E. (T.) candicans, E. (T.) thelegonoides, and E.leucantha.

N Chile(Region de Coquimbo northwards). This part of Chilecomprises two types of habitat, the Coastal Fog Desertwith irregular winter rainfall and the montane desert farinland with summer rainfall, the two being separated by

E the essentially lifeless Atacama Desert.

7%2 Coastal Fog Desert. This is very important in terms ofi endemic taxa, with more than 65 species, and it was the

subject of a conservation field survey sponsored by WWF-US (Anderson et al. 1990). Amongst the Cactaceae (ibid.:18-19; Hunt 1992) the remarkable genus Copiapoa, withabout 25 species, is endemic, and the treelike genusEulychnia (6 spp.) is almost endemic (1 sp. in S Peru).Other cactus taxa with endemic species are Eriosycc(Neoporteria), with 18 spp. restricted to the region(Kattermann 1994) Echinopsis (Trichocereus) (6 spp.),and Haageocereus (1 sp.), and there is also thetaxonomically isolated Opuntia miquelii. Representativesof o ther fami l ies a re Montiopsis and Cistanthe(Calandrinia pro parte, Portulacaceae) with 10 or more

a

Copiapoa desertorum plants uprooted, Chile.

116

Discocactus zehntneri ssp. boomianus, CITESAppendix I cactus known from only threepopulations.

mostly endemic spp., Deuterocohnia chrysantha(Bromeliaceae, endemic), Euphorbia ZactifZua, Oxaliscarnosa, 0. gigantea, and 0. succulenta, and various highlysucculent Nolana spp. Vegetation in this region is verypatchy and is represented by small green islandsinterspersed with extensive barren areas, their distributiondepending on how much moisture deposition from the fog(locally termed ‘Camanchaca’) takes place, which in turndepends on local land relief, height and steepness ofcoastal bluffs, and direction of the deep dry valleys(‘quebradas’). Almost every one of these ‘quebradas’ hasa distinct and characteristic vegetation, with someendemic species probably restricted to one ‘quebrada’only. The most up to date, comprehensive study aboutphytogeography, climatology, and ecology of this regionwas published by Rundel et al. (1991). However, the floraof the Coastal Fog Desert, known to be extremely rich inendemic species, remains poorly studied.

Hoffmann and Flores (1989) and Hunt (1992) assignIUCN conservation status categories to the aboveCactaceae and Bromeliaceae, but these werereinterpreted by Anderson et al. (1990) and in general thetaxa, though many are Rare, are not considered to beseriously under threat in view of the remoteness of thearea and generally limited human influence. However,there are important exceptions to this, notably as aconsequence of mining and associated ore treatmentactivities, goat grazing, rubbish dumping, local harvestingas timber, and commercial and amateur collection for thesucculent plant hobby. The species affected, all of themcacti and regarded as Vulnerable or Rare, are as follows:Copiapoa Maui (Rare, one of the smallest of all cacti andtherefore intensively searched for by collectors nearEsmeralda); C. megarhiza (Rare and insufficiently known,

but certaiat Paipotfrom ore

nly Ende, nearprocess

angered in theCopiap6, dueing); C. rupestl

lotto

*is (

ality surveyed in 1990very heavy pollutionRare, and apparently

now extinguished at its northernmost site, north of Taltal,by collecting for the horticultural trade); Eriosyce lmi(Vulnerable, from mining subsidence destroying itsmontane habitat south of Tocopilla); E. (Neoportcl-ia)napina (Vulnerable, from rubbish dumping and urbanspread in the vicinity of Huasco); E. occulta (Vulnerableand now apparently extinct through over-collection at thetype locality near Taltal, the more southern of its twoknown sites); E. heinrichiana (‘N. jussieui’) and E. sedisssp. elquiensis (‘N. nidus’) (Rare/Vulnerable throughovergrazing by goats in the region of La Sercna);Edychnia iquiquensis (Vulnerable - all populations knownin northern Chile are dying and there is no regenerationperhaps due to climatic changes leading to greateraridity); Edychnia sp. indet. (Vulnerable, PaposoTaltalregion, where affected by acid fumes from ore processingand local extraction for timber). El-iosyce manta (E.sandillon), a widespread Chilean endemic ranging intoCentral Chile, is sometimes collected for planting in localgardens and may be vu lnerabletowns, e.g. in the La Serena

wherearea.

it occurs near largeEchirqsis glauc~,

Eulychnia aricensis, and Haageocereus arrstdis arepossibly Rare/Vulnerable but insufficiently known taxa,

Uebelmannia pectinifera ssp. flavispina, MinasGerais, Brazil. CITES Appendix I.

117

which may not be regenerating due to the same climaticchanges as are affecting Eulychnia iquiq~~~is.

This part of Chile includes the established Pan deAzucar and Fray Jorge National Parks (WCMC 1992) andanother of considerable size under consideration since1985 in the region of Paposo (north of Taltal). Theseareas could protect a considerable number of thesucculents found in the region, and the more so ifrecommendations of the Anderson et al. (1990) report forthe modest enlargement of the more northern parks arecarried out. This would give protection to additionalspecies currently just outside park limits (amongst thesethe Rare Copiapoa rupestris and C. desertorum from southof Taltal, a second population of the rare C. Maui, and twoCopiapoa sp. nov. from Quebrada Botija, 70 km north ofPaposo). However, one problem to be resolved in the Pande Azucar National Park is the increasing population ofGuanacos (from 17 to >300 in the period 1983-1990)which graze upon the tuberous rootstocks of thegeophytic cacti.

lMontane Desert. Phytogeographically the Montane Desertis almost completely separated from the Coastal FogDesert. The cacti there (no other succulents are knownfor this region) have affinities to those from the adjoiningAndean countries. At lower elevations (between c. 2000-3000 m) there seems to be no regeneration of any cactuspopulation at present, which may be due to climaticchanges. The conservation status of the cacti from theregion has been assessed by Hoffmann and Flores (1989).The vulnerable taxa they identify are: Browningiacandelaris, Oreocereus (Arequipa) australis (endemic), 0.(A.) hempeliana, Haageocereus fascicularis (endemic),Neowerdermannia chilensis, Echinopsis atacamensis(endemic, but perhaps conspecific with E. pasacana fromArgentina), E. uebelmannianai (endemic), and Opuntiaconoidea (endemic). The taxonomic and thereforeconservation status of all Opuntia taxa from the regionremains unclear. All cactus populations from the higherelevations (>3000 m) seem to be out of danger.Corryocactus brevistylus and Neowerdermannia chilensis,assessed by Hoffmann and Flores as Vulnerable, arewidespread and there is no evidence that N. chilensis iscollected for human consumption as occurs with N.vorwerkii in Bolivia. In this part of Chile the LaucaNational Park and some other Reserves have beenestablished (WCMC 1992) but these include none of thethreatened species.

The Brazilian Region

This vas t r eg ion i s fu r ther d iv i s ib le on acountry/vegetational basis as follows:

Extra-Amazonian Brazil and Easternmost BoliviaFour major vegetational areas are recognised here, inorder of their importance for succulent plants. The largestsucculent genera, each with 15 or more endemic species,

Melocactus pachyacanthus ssp. viridis, known onlyfrom two sites surrounded by agricultural fields,northern Bahia, Brazil. Endangered.

are Dyckia, Encholirium, Pilosocereus, Rhipsalis, andMelocactus (Smith and Downs 1974; Zappi 1994;Barthlott and Taylor 1995; Taylor 1991b). Only recentlyhas the importance and plight of Brazilian succulentsbeen specifically addressed by that country’s authorities,with the proposal to place various endangered cacti onAppendix I of CITES. However, the most seriousproblem at present is the almost complete lack of reservesto protect the many rare and endemic terrestrial (ratherthan epiphytic) succulents from the dry parts of north-eastand south-east Brazil, and from the rocky East BrazilianHighlands, which rise out of the dry zone.

Eastern Brazil. This area includes the seasonally dry,deciduous thorn forests (‘caatinga’ and ‘agreste’) andassociated highlands (‘campos rupestres’) of north-eastern Brazil, plus rock outcrops in the savannas(‘cerrados’) of the central/eastern parts of the adjoiningstates of Tocantins and Goias, and, going south intosouth-eastern Brazil, dry areas and campos rupestres inthe states of Minas Gerais (excluding the extreme westand south-west) and Espirito Santo (inland valleys andinselbergs only). This area is home to about 100 broadlydefined Cactaceae species (90 per cent endemic, including10 endemic genera, Taylor and Zappi, ined.), c. 80 morenarrowly defined, 2 succulent species of Bromeliaceae,represented by Encholirium (endemic, c. 20 spp.) andnumerous species of Dyckia (c. 90 per cent endemic), andmany, mostly weedy and inadequately understoodPortulaca and Talinum taxa, besides a few succulents fromother families.

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Of special conservation concern are endemic speciesof the cactus genera Discocactus (6 spp., 5 endemic),Uebelmannia (endemic, 3 spp.), and Melocactus (M.conoideus, M. deinacanthus, M. glaucescens, M.paucispinus), which are Crit ically Endangered,Endangered, or Vulnerable and have been placed onAppendix I of CITES to afford them protection from theexport trade. Most of these endemics are known fromonly one or very few localities, where the populationsnumber between less than ten to at most 500 individuals.Discocactus horstii, D. placentiformis, D. pseudoinsignis, D.zehntneri ssp. boomianus, Melocactus glaucescens, M.paucispinus (both known from only two or three smallsites each), and all Uebelmannia spp. (U. buiningiiCritically Endangered, cf. Braun and Esteves Pereira1988) are threatened primarily by trade, including regularcollection of plants for seed production, or of seeds inhabitat for wholesale export in large quantities.Discocactus bahiensis and Melocactus deinacanthus (thelatter with only two populations known) are moreseriously threatened by agricultural development, andboth the former and D. zehntneri ssp. zehntneri had theirranges and numbers significantly reduced by inundationfrom the Represa de Sobradinho, a huge dam lakecreated in the 1970s on the Sao Francisco River(Bahia/Pernambuco). Repeated commercial collectingwas only partly responsible for the decline of Melocactusconoideus, a species Critically Endangered due to theextraction of the quartz gravel in which it grows, andthreatened with extinction at its type and only knownlocality above the expanding city of Vitoria da Conquista,southern Bahia (Taylor 1992). Some documented plantsstill exist in cultivation and could be used to effect itsreintroduction to the wild, should attempts to find newpopulations near its original habitat fail. The tallcolumnar species, Micranthocereus dolichospermaticus(from karstic Bambui limestone outcrops of difficultaccess in south-west Bahia), has attractive young seedlingsappreciated by the horticultural trade and may be indanger from the practice of felling mature individuals tofacilitate the collection of seed for wholesale export.Export of seed is not controlled for CITES Appendix IIspecies such as this, which deserves further investigationin habitat to determine if it should be proposed forAppendix I listing.

Threats

The driest zone of eastern Brazil, namely the ‘caatinga’and its ecotones with Atlantic Forest to the east (knownas ‘agreste’), dry forests to the south (in Minas Gerais andEspirito Santo) and savannas (‘cerrados’) to its west(Maranhao to Goias), represents a severely disturbedecosystem (Andrade-Lima 1981), which has been subjectto forest clearance for agriculture over more than twocenturies. However, in general many succulents haveprobably suffered less than most other plants as aconsequence of their frequent occurrence on rock

Esposfoopsis dybowskii, Bahia, Marack, Brazil.

outcrops unsuitable for cultivation or livestock grazing.Thus, many species of succulent Bromeliaceae (Dy&ia,Encholirium), Coleocephalocereus, various Pilosocereus,and some Melocactus (e.g. 111. ernestii, M. oreas) havesignificant populations in places dominated bygneiss/granite inselbergs, which are probably at less riskfrom habitat modification unless situated near expandingtowns. Of those cacti that are not mainly restricted to rockoutcrops, the least threatened are those which seem ableto regenerate when their forest habitat is cut over. Theseinclude Cereus jamacaru, Pereskia grandifolia, P. bahiensis,and P. stenantha, and all are also conserved by their USC: inthe form of impenetrable livestock fences or as hedgessurrounding homesteads, both within and sometimesoutside their natural ranges. A few very widely distributedendemic cacti which inhabit little-utilised or sufficientlydiverse habitats are probably not at risk, even thoughtheir numbers may have dropped significantly, e.g.Facheiroa squamosa, Harrisia adscendens, Opun tiainamoena, Pilosocereus gounellei ssp. gounellei and P.pachycladus s.1. However, other, mostly wide-rangingsucculents that are mainly found growing in the soil of thecaatinga-agreste, or on exposed rocks more or less levelwith the floor of the surrounding thorn forest, havesuffered considerable reductions in their distributions andabundance through forest clearance. Endemic cactusspecies affected in this way, whose ranges now appear tobe strongly fragmented, include Arrojadoa penicillata, A.rhodan tha, Brasilicereus phaeacan thus, Cereus albicarrlis,

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Coleocephalocereus goebelianus, Melocactus salvadorensis,M. zehntneri, Opuntia palmadora, Pereskia aureiflora,Pseudoacan thocereus brasiliensis, Pilosocereus ca tingicolas.l., P. J?occosus ssp. quadricostatus, P. flavipulvinatus, P.glaucochrous, P. pentaedrophorus s.l., Stephanocereusleucostele, Tacinga braunii, and T. funalis. Although mostof these are unlikely to become seriously threatened inthe immediate future, regular monitoring is essential ifsome are not to become endangered in the longer term.The same applies to some locally abundant andspectacular caatinga bottle-trees or ‘barrigudas’ from theBombacaceae (Cavanillesia arborea, Ceiba insignis s.1, C.jasminodora and Ceiba sp. indet. [SW Bahia]), whosehabitats have decreased sharply, especially in southernBahia, adjacent Minas Gerais and drier parts of westernEspirito Santo.

Of more urgent concern are Melocactus azureus ssp.azureus and M. pachyacanthus, which have smaller rangesand are restricted to local low-lying outcrops of limestone,whose vegetation gets destroyed when the surroundingcaatinga forest is cleared for cultivation. These taxashould be classified as Endangered on the basis of theirknown populations (see Taylor 1991b: 40-41), but furtherfield studies are needed in the remoter parts of northernBahia state, where additional and less disturbed habitatscould exist.

Other succulents from the caatinga, whose nativepopulations may be threatened, include the complex ofspecies allied with Euphorbia phosphorea, certainmembers of the Cucurbitaceous genus Apodanthera (e.g.A. succulenta, A. congestiflora), Dioscorea basiclavicaulis,and Marsdenia sessilifolia, but, unfortunately, little isknown about their conservation status, although someappear to be Rare or of restricted distribution (Rizzini1989, Jeffrey 1992, Rizzini and Mattos-Filho 1992). Evenif succulents found on raised rock outcrops within thecaatinga are generally at less risk from agriculturaldevelopment etc., some, and particularly those close toroads or human settlements, are at risk from thequarrying of stone for building materials. Those foundonly on limestone outcrops are probably most at risk (viz.Encholirium [3 spp. indet. cited by Andrade-Lima 1977:1911, Facheiroa cephaliomelana s.l., Melocactus azureusssp. ferreophilus, M. levitestatus, Micranthocereusdolichospermaticus, M. estevesii, Pilosocereus albisummus,P. densiareola tus, P. diersianus, P. flexibilispinus, P.~ZOCCOSUS, P. gounellei ssp. zehntneri, Opuntia saxatilis, 0.estevesii), but gneiss, granite, and other crystalline rocksare also quarried and, if this should take place at thesite(s) of one of the very local taxa, extinction could besudden (e.g. Marsdenia megalantha [Mun. Iramaia, BA],E n c h o l i r i u m sp . nov . [Mun. Tanhacu, B A ] ,Coleocephalocereus purpureus, Espostoopsis dybowskii,Melocactus deinacanthus, Opuntia werneri). 0. werneri isalready threatened at one of its localities through granitequarrying (Rui Barbosa, BA) and the other species areeach known from only one or two localities.

The few and mostly relatively small protected areaswithin the vast caatinga zone are as follows:

l Parques Nacionais Serra de Capivara (includesPilosocereus piauhyensis) and Sete Cidades (both inPiaui state),

l Estacao Ecologica de Serido (Rio Grande do Norte),l Reserva Ecologica Raso da Catarina (NE Bahia),l Reserva Biologica Federal da Serra Negra

(Pernambuco),l Areas de protecao ambiental da Serra de Baturite

(Ceara) and Gruta dos Brejies / Vereda do RomaoGramacho (Bahia, includes Melocactus azureus),

l Estacao Ecologica Federal de Aiuaba (Ceara).

These can offer protection to only few and mostly thewidespread species noted above, since, unfortunately,there are currently no significant protected areas in thesouthern part of the caatingas zone (central-S Bahia to NMinas Gerais), where higher species diversity andendemism is matched by a most disturbing level of habitatdestruction (mainly for agriculture and charcoalproduction). One of the most important areas needingprotection amongst the southern caatinga-agrestes is themiddle section of the Rio Jequitinhonha valley (i.e.Aracuai to Jacinto) in north-eastern Minas Gerais, wherea remarkably rich assortment of succulent plants exists(Rizzini and Mattos-Filho 1992) including many endemicand potentially threatened cactus species (Taylor andZappi 1992). Another promising site for protection, with acomprehensive range of southern caatinga cacti, includingthe rare Espostoopsis dybowskii, is situated to the east ofthe village of Porto Alegre, on the north bank of the Riode Contas, Mun. Maracas, Bahia. Other sites need to beidentified for the conservation of succulent taxacharacteristic of the deep soils and Bambui limestoneoutcrops in the valley of the Sao Francisco River(especially for Bombacaceae and columnar Cactaceae).One such would be the massive raised outcrop south ofthe town of Iuia on the east bank of the river (Bahia),which, besides some spectacular bottle-trees ofCavanillesia and Ceiba growing around its base, has twovery local endemics (Facheiroa estevesii, Opuntia estevesii)restricted to the rock itself. Other sites should be foundon the west side of the river, where further endemics, suchas the aforementioned Micran thocereusdolichospermaticus and Facheiroa cephaliomelana, arelocated.

The East Brazilian Highlands, with their mosaic of‘camp0 rupestre’ and ‘cerrado’ vegetation (Giulietti andPirani 1988; Zappi and Taylor 1994: 77) represent theleast modified of the environments considered under thepresent heading of eastern Brazil. However, they have asmany if not more endemic succulent species than thecaatingas-agrestes just discussed, and many are ofextremely local occurrence and therefore potentially atrisk. Widespread and mostly common, non-threatened

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exceptions include Cottendorfia florida, Cipocereusminensis ssp. minensis, Leocereus bahiensis, Melocactusbahiensis, M. concinnus, Pilosocereus aurisetus ssp.aurisetus, Micranthocereus purpureus, and StephanocereusZuetzelburgii, the latter two endemic to the extensiveuplands of the Chapada Diamantina, Bahia, and alsofound within its national park (Mucuge-Lencois).

Utilisation of the campos rupestres is limited to cattlegrazing, with associated burning to induce regrowth, andlocal extraction of some plants, e.g. Eriocaulaceae (driedflower export trade - a serious conservation issue),orchids, and VeZZozia spp., and there is also limiteddisturbance caused by small scale mining for gold andprecious stones. Some parts where cerrado vegetation ismore abundant are being cut over for the production ofcharcoal and later converted into Eucalyptus plantations,especially in Minas Gerais state, where this activity is oneof the factors threatening UebeZmannia spp. andCipocereus crassisepalus. The burning for cattle grazingdoes affect some native populations of succulents, but theregular collection of plants, and nowadays more especiallyof seed? of certain rare cacti may be cause for greaterconcern.

In addition to some of the CITES Appendix I taxanoted above, the following campo rupestre / cerrado cactiare known from only one or two small populations, or atbest have a very localised range which does not includeany kind of designated protected area (cf. Taylor andZappi, ined.): Arrojadoa dinae (especially the rare variantknown as A. eriocaulis), Arthrocereus rondonianus,Brasilicereus markgrafii, Cipocereus bradei, C. crassisepalus,C. laniflorus (sp. nov. ined.), C. pusilliflorus, Melocactusviolaceus ssp. ritteri, Micranthocereus albicephalus, M.auriazureus, M. polyanthus, M. streckeri, M. violaciflorus,Pilosocereus vilaboensis, P. aurisetus ssp. aurilanatus, andP. fulvilanatus ssp. fidvilanatus and rosae.

Similarly restricted taxa located within protected areasare rather few: Arrojadoa bahiensis (Parque NationalChapada Diamantina, Bahia), Cipocereus minensis ssp.pEeurocarpus (Parque National da Serra do Cipo, MinasGerais), Arthrocereus melanurus ssp. nov. (ParqueEstadual de Ibitipoca, MG) and Pilosocereus rupicola(Estacao Ecologica da Serra de Itabaiana, Sergipe). Ifextended slightly to its west, the first-listed would includea second population of the remarkable A. bahiensis. Thelast-named Pilosocereus is possibly endangered or evenextinct, but has not been investigated in habitat in recenttimes. The Serra da Piedade (Mun. Caete, MG) is not adesignated protected area, but a site of religioussignificance, which has a population of Arthrocereusglaziovii, a specialised species restricted to rocks very richin iron (‘canga’), many of its former habitats havingdisappeared through ore extraction. It is also the type andonly known locality for Dyckia simuZans. Numerous, other,little-known species of Dyckia and some Enchohrium arerecorded from various serras in central and eastern Goiasand especially from the regions of Diamantina, the Serrado Cipo and serras further south in Minas Gerais state.

Melocactus viokceus, on the coast of Bahia, Brazil,threatened by tourist developments.

These deserve further study in the field to determine theirtaxonomic and conservation status. A peculiar andspecialised cactus, found in the sandy cerrados borderingon the caatinga and campo rupestre zones, from westernBahia to central-eastern Minas Gerais, is Cc~rrsmirabella. It is widespread but of erratic occurrence andmuch of its habitat is being destroyed by charcoalproducers, so its status needs to be monitored carefully.

Locations where new protected areas are needed toassist the conservation of the above listed rarities,including the earlier discussed CITES Appendix I cactustaxa, are as follows and are listed in the Action Proposals.

Atlantic Forest. This comprises the coastal rain forest(‘Mata Atlantica’ in its strictest sense) and sandy littoraldunes (‘restingas’) of north-eastern Brazil and theirextensions southwards, where the former broadens andmerges with the planalto forests of south-eastern andsouthern Brazil, reaching the northern part of RioGrande do Sul state. The area, which is very humid, has ahigh diversity of epiphytic cacti from the tribeRhipsalideae, but, as is now well known, only a smallfraction of the original forest remains. EndemicRhipsalideae include the horticulturally and economicallyimportant genera Schlumbergera (6 spp.) and Hatiora (5spp.). A few very widespread or regionally common taxa,such as the epiphytic H a t i o r a salicornioides f.salicornioides, Lepismium cruciforme, L. houlletianum, L.warmingianum, Rhipsalis floccosa, R. lindbergiana,R. elliptica, and R. cereuscula, and the non-ep

R.‘P

teres,hvtes

Brasiliopuntia brasiliensis, Opuntia monacantha, Cereusfemambucensis, Pilosocereus arrabidae, and P. brasiliensisare probably to be regarded as at low ris k, but theremaining Brazilian endemic species are of conservationconcern to varying degrees. For example, the wide-ranging but erratically occurring restinga taxa, Melocactusviolaceus ssp. violaceus and margaritaceus, are threatenedat various points in their ranges by touristic developmentsand other forms ofdiversity ‘hot spots’

urbanare fou

expansion. Partitular speciesnd in southern Espirito Santo

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state (between Domingos Martins and the Serra doCaparao) and around and between the great cities of Riode Janeiro and Sao Paulo, various taxa being endemic tovery small areas. The flora of southern Espirito Santo ispoorly understood, but includes a recently describedspecies of Christmas Cactus, Schlumbergera kautskyi(known from only two or three small sites), and theremarkable, red flowered Rhipsalis hoeZZeri. Other speciesare represented by disjunct populations, often at theirnorthern limits, such as Hatiora sahcornioides f. cylindrica,Rhipsalis cereoides, R. pilocarpa, R. campos-portoana, andSchlumbergera microsphaerica (the latter two within theboundaries of the Parque National do Caparao).

Further south the diversity of epiphytic and epilithicRhipsalideae increases markedly along the coast and inthe Serra do Mar westwards from Cabo Frio (RJ), wherealso an isolated member of tribe Cereeae, Pilosocereusulei, is narrowly endemic on coastal rocks. Rhipsalispentaptera, a species relatively common in cultivation, ispresumed to be extinct in the wild, since its only recordednative site is within what is now the city of Rio de Janeiro(at Praia da Gavea). Other rare and probably vulnerable(but inadequately studied) species of Rhipsalideae fromthe region of Rio de Janeiro include Rhipsalis pacheco-leonis, R. cereoides, R. mesembryanthemoides, and

Schlumbergera orssichiana. Rhipsalis burchellii is knownfor certain only from the metropolitan region of S5oPaulo, and a substantial part of its presumed formerhabitat appears to have been either destroyed completelyor severely affected by industrial pollution. Hatioraherminiae and H. epiphylloides are each known from onlytwo relatively small areas of montane cloud forestbetween Rio and Sao Paulo and should be classified asendangered due to forest clearance, even though they areat least partly found within protected areas (the former inthe Parque Estadual Campos do Jordao, SP, the latter inthe Parques Nacionais do Itatiaia, RJ/MG, and Serra daBocaina, RJ/SP). Other species of Rhipsalideae from thispart of the Atlantic Forest and montane forest zonesappear to have greater ranges, but are infrequent, disjunctor seldom observed. These include Hatiora salicornioidesf. cylindrica, Rhipsalis neves-armondii, R. grandifloru, R.pilocarpa, R. clavata, R. pulchra, Schlumbergeru truncutu,S. russelliana, and S. opuntioides. Protected areas thatinclude or probably include one or other of these speciesare the Parques Nacionais da Floresta da Tijuca and daSerra dos Orgaos (RJ), the Parques Estaduais de IlhaGrande (RJ), de Ibitipoca (MG), de Campos do Jordaoand de Picinguaba (SP), and the Reservas Biologicas dePoco das Antas, de Paranapiacaba, da Jureia, Ilhabelaand that proposed for the Serra do Japi (SP).

Destruction of the Atlantic Forest has been greatest innorth-eastern Brazil, where only 5-10 per cent remainsand, therefore, our knowledge of the flora iscorrespondingly fragmentary. It is quite possible thatepiphytic Cactaceae from here have become extinctbefore discovery and description. In Paraiba andPernambuco remnants of this forest include the ‘brejos’on higher land away from the coast, where the watershedsare important for the human populations living belowthem. Such forests are currently being studied andcatalogued as part of an Anglo-Brazilian initiative(Plantas do Nordeste), with great emphasis being placedon the need to preserve these floristic refuges which, interaha, include disjunct populations of cactus epiphytes, suchas Lepismium cruciforme and Rhipsalis crispatu.

Further south, in coastal Bahia (up to 100 km inland),between the capital Salvador and Belmonte to the south,where annual rainfall is generally in excess of 1750 mm,there are occasional records of various species ofRhipsalideae, indicating a once rich centre of diversityincluding Hatiora salicornioides f. cylindricu, Rhipsulisparadoxa ssp. septentrionalis , R. bucciferu ssp.hileiabaiana., R. russellii, and R. oblonga. With so littleforest remaining it seems reasonable to assume that hereall of these are threatened to a greater or lesser extent,

*r 1 I-‘. P . ,- . 1 1~ even ir some may benerit rrom protection in local

$5 reserves, such as the Reserva Biologica Federal de Una2 (south of Ilheus).

Also part of the Brazilian north-east, is the

Rhipsalis pentaptera, almost extinct in the wild at the Archipelago of Fernando de Noronha, a Federal

only known site within the city of Rio de Janeiro. Environment Protection area. These Atlantic islands are

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home to at least one endemic cactus, Cereus insularis (arelative of the Brazilian coastal C. fernambucensis), whichseems adequately protected at present. A second species,or perhaps a form of the first, is C. ridleii, which has notbeen seen since its original collection in the 1950s andmay now be extinct. It is no longer in cultivation, so far asis known.

Some of the best-preserved Atlantic Forest andcoastal habitats are those found in southern Brazil, in thestates of Parana, Santa Catarina, and northern RioGrande do Sul, between sea level and almost 2000 m.Here a wide variety of Rhipsalideae could be protected ifdeforestation can be controlled. These include thehorticulturally important Easter Cacti, Hatiora gaertneriand LL rosea (its range includes Parque National de SaoJoaquim, SC), which are characteristic of Araucaria forest,the peculiar Rhipsalis dissimilis (a widespread butinfrequent SE Brazilian lithophyte protected in theParque Estadual Vila Velha, Parana), and the forestepiphytes Rhipsalis trigona, R. paradoxa, R. pulvinigera, R.puniceodiscus, R. pachyptera, and R. campos-portoana.There are also at least 11 endemic and 5 non-endemicDyckia spp., all of unknown conservation status. In thewestern part of southern Brazil, in the drainage of the RioParana, the situation is rather different, much of thehumid and savanna forest having been cleared foragriculture. Fortunately, there are no endemic succulentsknown from this vegetation, which extends westwards intoeastern Paraguay, where it is better preserved.

Besides those mentioned already, protected areas inthe species-rich Atlantic slopes of southern Brazil includethe Parque National de Superagui and the FederalEnvironmental Protection Area and Ecological Station ofGuaraquecaba (Parana), which probably include at leastsome epiphytic taxa. Non-endemic succulents found in theforested and coastal parts of southern Brazil that areprobably out of danger include Cereus hildmannianus andLepismium lumbricoides.

The third and fourth areas of extra-amazonian Brazilcomprise the plant communities mainly composed ofgrasses etc. (‘campos’) of southernmost Brazil (RioGrande do Sul state) and the savannas or ‘cerrados’ ofCentral-western Brazil. The relatively smaller succulentfloras of these two areas and the perceived threats havemuch in common with those of eastern Paraguay, easternBolivia, and Uruguay, as discussed below.

Campos. The ‘campos’ of Rio Grande do Sul areimportant for the high number of endemic and non-endemic but probably threatened taxa belonging to theCactaceae-Notocacteae, i.e. the genera Parod ia(Notocactus) and Frailea, and tribe Trichocereeae(Gymnocalycium). There are also two endemic species ofDyckia (Bromeliaceae), their conservation statusUnknown. Other succulents are representatives ofwidespread elements characteristic of the floras ofArgentina and Paraguay and probably not at serious risk

as species, e.g. Cereus aethiops, Echinopsis spp., Pereskianemorosa, and Lepismium lumbricoides (an epiphyte). Asexplained below, under Uruguay, the habitats of many ofthe endemic and threatened Notocacteae arc rockyoutcrops amongst agricultural land, much of the terrainbeing cultivated for arable crops or grazing pasture.Exceptions include the Parque National Aparados daSerra, situated at the northern border of the state withpart inside adjacent Santa Catarina, and including thehabitats of the endemic Parodia haselbergii and P .graessneri, which are presumed to be adequatelyprotected. The conservation status of most of theremaining 40 or so Parodia (Notocactus) taxa (many ofdoubtful taxonomic standing), c. 15 Frailea and 4Gymnocalycium spp. (2 endemic), which are concentratedin the southern part of Rio Grande do Sul, needs to bedetermined. However, some populations are known to bevery small and illegal collection to satisfy the demand fornovelties by hobbyists in Europe and elsewhere iscertainly taking place. There appear to be no otherofficially designated, protected areas includingNotocacteae/ Trichocereeae in Rio Grande do Sul.

Western cerrados. The ‘cerrado’ in the states of MatoGrosso, Mato Gross0 do Sul, Goias, the western parts ofsouth-eastern Brazil and easternmost Bolivia (SantaCruz) comprise open savanna woodlands on oligotrophic(strongly weathered and leached) soils and included rockyoutcrops, inselbergs and uplands, such as the Chapadados Guimaraes (Mato Grosso). The altitude variesbetween 300-1500 m and rainfall is in excess of 1000 mmper annum with high average temperatures for most ofthe year. This area has very few endemic succulents,including 8 poorly known species of Dyckia and only 5botanically distinct species of cacti: Arthroccreusspinosissimus, Cereus adelmarii, C. saddianus, Echinopsishammerschmidii, and Frailea chiquitana (the latter two areBolivian endemics found on inselbergs in the ecotonalregion between the ‘cerrado’ and Amazonian forests), allof whose conservation status is inadequately known (theArthrocereus may benefit from any protection afforded byits location inside the Parque National da Chapada dosGuimaraes). Widespread cactus taxa include the highlyvariable Discocactus heptacanthus s.1. and Pilosocereusmachrisii s.1. (both fragmented into numerous ill-definedmicrospecies by some authors), besides a treelike speciesof Cereus found on calcareous outcrops, whose identity isuncertain at present (C. calcirupicola ?), C. hicolor(botanical affinity uncertain) and the shrubby C. euchlorus(= Praecereus sp.) . One o r more Opuntia spp.,Cleistocactus horstii (? = C. baumannii), Frailcacataphracta s.1. (including F. matoana, EW according toH u n t 1992)) C e r e u s ( M o n v i l l e a ) kroenleinii,Gymnocalycium anisitsii, G. marsoneri (G. matoensc),Harrisia guelichii (Cereus balansae), Pereskia sa~~haros~,Jacaratia corumbensis, Dyckia @ox, D. microcalyx, andDeutercohnia meziana also occur, but have the major parts

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of their ranges in adjacent Paraguay. Although thefireswept ‘cerrados’ are not noted for succulent plants,there may be reason for concern about the status of eventhe widespread taxa in this large geographical area, andespecially taxa restricted to calcareous soils and rocks,which are at greater risk from habitat conversion. Largescale farming operations, for both arable (especiallysoybean) and livestock, are modifying the environmentand local assessments of the likely effects on succulentplant populations are needed. In easternmost Bolivia thisenvironment has been used for cattle grazing for morethan 200 years and is now being cultivated in some areas.The seasonally inundated region of the Pantanal (Brazil),which includes raised rocky areas where succulents arefound, is currently benefitting from ecotourism and maythus be less at risk from agricultural development.

E Paraguay and NE ArgentinaThis subdivision includes two of the three vegetationalareas of Paraguay (Esser 1982; Metzing 1994) namely thevalley of the Rio Paraguay and the part of Paraguay to itseast, and the north-eastern Provinces of Misiones andCorrientes from Argentina. The succulents of this areaare almost exclusively Cactaceae and Bromeliaceae(Dyckia spp.) and many of these are also found in theadjacent parts of central-western and southern Brazil (seeabove). According to Metzing (see also references toEsser 1984a, b cited therein) some of the cacti of easternParaguay are seriously affected by agricultural practices,including the destruction of protective nurse-plant shrubsby fire, and the same threats apply in north-easternArgentina, where forest clearance is more accentuated.Genera with one or more species affected in this wayinclude Cereus, Opuntia, Pereskia, Gymnocalycium, andHarrisia, some of which include endemics whoseconservation status needs further investigation (e.g. G.mesopotamicum, H. hahniana, the first-named knownfrom a single, flat, rock outcrop where grazing occurs).More specific problems are noted below.

Between 30 and 35 species of Cactaceae are knownfrom this region, of which the following are endemic toParaguay and have been assigned regional conservationstatus categories by Metzing (1994) who believes theydeserve to be included on Appendix I of CITES: FraiZeaknippeliana (Rare, only 2 localities), Gymnocalyciumparaguayense (Endangered, only 2 small populations,which could be quickly eliminated by collecting and arecurrently affected by livestock grazing); ‘G.fleischerianum’ (Endangered, 2 5 localities known,affected by tourism, house construction etc.) and Parodia(Notocactus) nigrispina (Endangered, only 3 populations,affected by land clearance, grazing and collected for saleat the roadside). The last two are restricted to thecrystalline ‘Cordilleras’, where the peculiar endemic,Cereus lanosus (probably not threatened), is also found.Non-endemic species discussed by Metzing (1994)include: Frailea cataphracta (ranging to adjacent CW

Brazil and Bolivia, vulnerable), Parodia (Notocactus)ottonis (Vulnerable/ Endangered in Paraguay throughland clearance and grazing, but still common inneighbouring countries), P. (N.) schumanniana ( N EArgentina and S Brazil, Rare) and Pilosocereus machrisii(P. juaruensis) (Vulnerable in Paraguay, but widespreadand not threatened in adjacent Brazil). The non-endemicDiscocactus heptacanthus ssp. magnimammus (D.hartmannii), from the campos cerrados of north-easternParaguay, is potentially threatened from conversion of itshabitat for agriculture.

Besides cacti, there are 5 endemic species of Dyckiadescribed from eastern Paraguay (D. exserta, D.velloziifolia, D. affinis, D. tobatiensis, D. tomentella) and 3from Misiones Province, Argentina (D. niederleinii, D.subinermis, D. mitis), all known only from the type or veryfew collections (Smith and Downs 1974).

Of the protected areas listed in WCMC (1992) theBiological Reserve of Itabo includes the habitat of variousnon-endemic epiphytic cacti (D.C. Zappi, pers. comm.),while the Cerro Cora National Park may give someprotection to Brasiliopuntia brasiliensis, Pilosocereusmachrisii, and the endemic Dyckia exserta (Bromeliaceae).It is not known whether the forest reserve of Capivary(13,500 ha) includes and protects populations of the non-endemic Discocactus heptacanthus ssp. magnimammus (D.hartmannii), whose genus is on Appendix I of CITES.

SE Bolivia, W Paraguay and N ArgentinaThis is the arid vegetation type known as the Chaco,which is often compared with the Caatinga of north-eastern Brazil, but has, in fact, very few floristicsimilarities and a quite different succulent flora.Widespread and conspicuous cactus species includeQuiabentia verticilla ta, Cereus (Praecereus) saxicola(h4onvillea cavendishii misapplied), C. spegazzinii,Browningia (Castellanosia) caineana, Cleistocactusbaumannii, Gymnocalycium marsoneri, G. mihanovichii,G. pflanzii, Harrisia pomanensis, Opun tia quimilo, 0.retrorsa, Pereskia sacharosa, and Stetsonia coryne, none ofwhich is believed to be particularly threatened at present,though up-to-date knowledge is poor and environmentalchange in the form of clearance for agriculture isaccelerating. Another widespread succulent is Ceiba(Chorisia) speciosa (Bombacaceae).

SE Bolivia. The extensive low forest, spiny scrub and drysavanna loosely referred to as Chaco in south-easternBolivia is characterised by a hot climate with precipitationof 500-1000 mm/yr concentrated in the four warmestmonths. Little knowledge of the succulents found here iscurrently available, but habitat alteration appears to belimited. However, this could change rapidly if modernmachinery is employed on a large scale, as is beginning tooccur in Paraguay.

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WParaguay. Here Chaco vegetation is said to cover 60 percent of the territory of Paraguay, yet this area counts lessthan 100,000 in human population. However, it has agreater number of endemic species than in the adjacentChaco countries, e.g. Cereus lamprospermus, C.pachyrhizus, C. phatnospermus, and Gymnocalyciumeuqpleurum, and G. paediophilum (both rare and knownonly from or near Cerro Leon). According to a LandUtilisation Survey published in 1991 and based onsatellite images, only 4.22 per cent of the total area isunder cultivation. However, a somewhat differentimpression is gained from observations on the ground,where large cultivated fields can be seen along the Trans-Chaco highway (and especially in the vicinity ofFiladelfia). There is also selective extraction of timber andcattle ranching in some places, but much of the Chaco isinsufficiently explored to give an accurate assessment.According to Metzing (1994) Cerro Leon deserves to bedesignated a National Park. The vegetation in its vicinityis relatively undisturbed at present.

Further west the soils become more sandy and dunessometimes occur. Here the vegetation is more open,grasses and other herbs predominating, the shrubs inscattered groups (‘espartillares’). The land is potentiallysuitable for cultivation and pasture, which might lead tothe endangerment of the non-endemic Gymnocalyciummegatae. A distinct species said to come from Paraguayand perhaps from the Chaco region, but of unknown wildstatus today, is Cereus haageanus.

N Argentina. The Chaco environment of Argentina hasbeen modified on a much greater scale than that inParaguay. For many years timber of the dominant-emergen t quebracho t r ees (Sch inops i s s p p . ,Anacardiaceae) has been extracted as raw material fortanning, manufacture of railway sleepers, and forfirewood. The remaining stumps produce sucker sprouts,which are eaten by cattle, preventing regeneration, andthe remainder of the vegetation is often exploited forcharcoal. Thereafter, the land becomes pasture or is leftabandoned, but does not seem able to return to theoriginal climax vegetation. Gymnocalycium spp. sufferimmediately after the forest cover is removed, but othersucculents, such as Quiabentia verticillata and Frailea spp.,seem to withstand habitat modification and may evenincrease in abundance in some areas.

The Chile-Patagonian Region

The cactus genera Austrocactus (6 spp.) and Maihuenia (2spp.) are endemic to this region. Another importantcactus genus, almost endemic to the region, is Pterocactus,an isolated taxon of Opuntioideae with 5 species endemicto Argentinian Patagonia. P. australis is said by Kiesling(1990) to be the most southerly occurring species ofCactaceae.

UraguayDiscounting a few, mostly weedy species of Portulaca andthe non-endemic and wide-ranging Dyckia remotiflora(status unknown), the native succulent plants of Uruguayare all Cactaceae, the most important belonging to thetaxonomically complex, dwarf, globular genera Parodia(incl. Notocactus), Gymnocalycium, and Frailea. Thesethree comprise more than 50 species in total, the majorityof which, though only provisionally accepted (Hunt 1992)are said to be endemic. Even if many of these taxa are“less-than-species”, their conservation status is cause forserious concern, since some 75 per cent of the landsurface of Uruguay is said to have been modified withpositive or negative effects by agriculture etc. (mainlycattle grazing) and there are scarcely any protected areas(WCMC 1992). Populations of such cacti are often verysmall, being restricted to isolated rock outcropssurrounded by pasture. Various of the non-endemicglobular taxa are otherwise known only from theneighbouring Brazilian state of Rio Grande do Sul (seeabove), where similar threats apply. Other cacti, such asspecies of Cereus, Cleistocactus, Harrisia, Opuntia,Pereskia, and epiphytic genera, i.e. about 20 species in all,have probably been severely reduced in abundance. Mostof these are widely distributed in the adjacent countries ofArgentina, Brazil, and eastern Paraguay, where similarproblems exist, though to a lesser degree. While reliableinformation is lacking it is impossible to make suggestionsfor action other than to instigate field surveys by botaniststaxonomically competent in the species-rich globulargenera noted above. There are also logistical difficultiesin accomplishing such work, since much of the habitat isprivately owned land and good contacts with local farmerswould therefore seem a prerequisite to any kind of fieldstudy.

ArgentinaThree geographical/vegetational subdivisions can berecognised here: the Monte, the Pampas, and Patagoniaproper.

Monte. This is the western desert of Argentina, to the eastof the high Andes, and characterised by a spiny scrub inwhich Larrea (Zygophyllaceae), the creosote bush,dominates. Agricultural activity occurs only in smallpatches near rivers, due to the irregular and limitedrainfall (50-300 mm/yr), but farming of goats and sheep issometimes intense, especially near villages or isolatedhouses, where disturbance of the natural vegetation isgreatest.

Three genera of Cactaceae are represented bynumerous species here: the endemic Tephrocactus Group(s. str.) of Opuntia, Gymnocalycium, and Echinopsis(Trichocereus). Some of the Tephrocactus species have apartly clonal mode of reproduction, and others are knownfrom single populations or very limited areas (e.g. 0.halophila, 0. molinensis), but none seems to be seriously

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ti

Rain sticks of Eulychnia and Echinopsis, La Serena,Chile.

threatened. The same applies to the numerousGymnocalycium spp., although G. schickendantzii plantscan be damaged by the trampling of livestock. Echinopsisstrigosa, E. candicans, and E. angelesiae are frequentplants. Other succulents are represented by PoutuZaca (>3 ephemeral but often abundant spp.) and Talinum (3tuberous spp.), which do not appear to be severelyaffected by grazing.

Pampas. A huge part of the territory of Argentina wasnaturally covered in herbaceous prairies (the Pampas),where rainfall is regular, the climate warm temperate andthe soil fertile. This area originally comprised theprovince of Buenos Aires and S Santa Fe (humid pampa),and Province La Pampa and S Cordoba (dry pampa), butit has long been occupied by agriculture, which hasartificially extended its limits northwards into much ofSanta Fe and Cordoba (where spiny forest or ‘Espinal’formally occurred). Besides agriculture, the giantconurbation of Buenos Aires (13 million pop.), occupiesthe central-eastern part of the area.

Cacti and other succulents were never abundant here,although the substantial modification of the environment

makes assessment of their original status difficult anddepends on what can be observed growing on abandonedfarms and beside railway tracks, and sometimes used inhedges or gardens. Such species include O p u n t i abqnaerensis (= 0. paraguayensis), 0. monacantha (notthreatened), Ceveus uruguayanus (‘C. peruvianus ’ hort.),Gymnocalycium schroederianum (with a disjunctdistribution on low rock outcrops), and G. pZatense.

Patagonia. The succulents of this huge area south of theRio Negro are chiefly cacti (Kiesling 1990) and theendemic succulent Composite, Duseniella patagojzica,none of which is considered to be particularly threatened.The cold, dry climate has limited land use to sheepfarming, which has altered the floristic composition,possibly favouring the less edible cacti and their spinyshrub nurse-plants. Cacti such as Maihuenia patagonicu,Opuntia (Maihueniopsis) darwinii, Austrocactus spp., andperhaps even Pterocactus spp. may now be more abundantthan prior to habitat modification. Other cacti are mostlywidespread species that are also out of danger.

Chile(Region de Valparaiso southwards). Central andsouthern Chile is home to approximately 16 species ofCactaceae (9 endemic), some of which are known to beunder considerable threat, and one may be extinct. Thefollowing information is based mainly on Hoffmann andFlores (1989) and Kattermann (1994).

Straddling the border between the regions ofCoquimbo and Valparaiso (but counted amongst thelatter’s endemics here) are the Vulnerable Echinopsislitoralis and endangered Eriosyce chilensis, the latterknown from only an 11 km stretch of coastal rocksbetween Pichidangui and Punta Molles and severelythreatened by housing construction. Generally, tourismon the Chilean coast is increasing rapidly hand in handwith building projects for the leisure industry, especiallybetween San Antonio and Coquimbo, an area withvarious endemic Eriosyce species. According toKattermann (1994) Eriosyce aspillugae is probably extinctat its only known locality at Hacienda Tanume(Colchagua) due to habitat modification for pulpwoodproduction and collecting activities for horticulture(another population from Cabo Carranza, Talca Province,may possibly represent this species, but is currently ofuncertain identity - Kattermann, l.c. 73). However, thespecies is still extant as habitat-documented material in afew specialist hobbyist and botanical living collectionsoutside of Chile and in the National Botanical Garden atVina, which could be utilised to propagate stock forpotential reintroduction to the wild. Populations of theendemic E. cuwispina var. tuberisuZcata, from the vicinityof Valparaiso, are now reduced to only a few plantssurrounded by apartment complexes, and numbers of thewide-ranging E. subgibbosa have been severely reduced inthe vicinity of Conception by collection for the local

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markets at Christmas time, when they are in flower. E.engleri is known from a single locality in mountainsbetween Santiago and Valparaiso. Austrocactus spiniflomsis a rare and perhaps vulnerable species known only from1500-2500 m altitude in the Andes immediately to the eastof the capital, Santiago. A. hibernus, from further south,at 2000 m in the Cordillera de1 Maule, is variouslyregarded as either Rare, Vulnerable or Endangered (cf.Hoffmann 1989; Hoffmann and Flores 1989; Hunt 1992).A third Chilean species A. philippii, is insufficientlyknown. The only known locality for A. patagonicus inChile, near Chile Chico, has been destroyed by theeruption of the volcano Hudson in 1991, but the speciesstill exists in neighbouring Argentina. The endemic, butcommon and widespread Echinopsis chiloensis andEulychnia acida are regarded as being out of danger.Wood of the Edychnia is used to make percussioninstruments (‘palos de agua’) and currently commands a

considerable price, the finished articles entering theexport trade. At present only dead wood is used (L.Faundez, Univ. de Chile, pers. comm. to R. Kraus) andthis presumably does not represent a conservationconcern. The almost endemic Maihueh poeppigii isregarded as out of danger at its relatively inaccessible,unpopulated sites in the Andes of southern Chile.However, its localities in the Central Valley to the northare threatened by overgrazing and expanding forestryplantations of Pinus radiata. The only coastal habitat ofthis species near Escuadron will soon be destroyed byurban expansion (0. Matthei, Univ. Conception, per-s.comm. to R. Kraus).

The following individuals reviewed this section and offeredvaluable criticisms and additional data which have been dulyincorporated: D.C. Zappi & R.M. Harley (Kew), M.B. Christiansen(Copenhagen) and D. Metzing (Bremen). D.C. Zappi kindlytranslated R. Kiesling’s contributions from Spanish.

Eulychnia fence, RioLimari, Chile.

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Chapter 4

Action Proposals

The actions required to conserve the world’s diversity ofsucculent plants are many and varied. The reviewundertaken by the IUCNSSC Cactus and SucculentSpecialist Group in order to prepare this documentindicates some of the main activities which should beundertaken immediately. These can be divided intovarious broad categories. Some are straightforward andrelatively inexpensive: for example, provision ofinformation to conservation decision makers. Others aremore complex, involving, for example, long-termdevelopment of protected area systems. As far as possibleorganisations have been identified for the actionproposals listed below. The Specialist Group will workwith these organisations to develop detailed fundingproposals for essential conservation action. The SSCGroup will also work with 10s to bring to the attention ofnational governments the need for effective conservationlegislation, the development of effective scientific andmanagement authorities to enforce national and CITESlegislation, the need for protected areas, and otherconservation initiatives to protect the world’s diversity ofsucculent plants. It is our hope that researchers andstudents, funding agencies, conservation organisations,specialist groups, societies, collectors and growers will allfind this document, and especially the Action Proposals,helpful while playing their part in plant conservation.

General1) Provision of information. This Action Planbrings together certain data for the first time.Information compiled for this report on the conservationstatus of species has been incorporated into the PlantsDatabase maintained at WCMC. Provision needs to bemade for the regular updating of species data by theIUCN/SSC Cactus and Succulent Specialist Group inassociation with WCMC. It is also a priority to assign thenew IUCN categories of threat to those species ofconservation concern indicated in this Action Plan.

It is particularly important that information on theconservation status of species coupled with data onpriority habitats for succulent species conservation shouldbe maintained and made available to national protectedarea agencies for use in designing protected area systems.

2) Development of proposals to amend theAppendices of CITES. Recent information, includingthe results of various field projects and nursery surveys,indicates that the following changes should be made tothe CITES Appendices. The SSC group will help therelevant national CITES management authorities todevelop proposals accordingly.

a>

b)

C)

3)

Addition of Beaucarnea to Appendix II of CITES. Allnine species in the genus are threatened, mainly dueto collection by the horticultural trade. Over-collection of seed and seedlings is also damaging wildpopulations. A proposal should be developed to addthe genus to Appendix II of CITES.

Contact: UAT and Dr. Luis S. Herrdndez

Addition of other taxa to Appendix II: A&FZ~II(Madagascan species), Adenium, Brachystelma,Cyphostemma, Commiphora (Madagascan species.),Fockea, Ha worthia, Kedrostris, Nolina, Odosicyos,Operculicarya, Raphionacme, Trochomeropsis,Xerosicyos, Zygosicyos.

Transfer from Appendix II to Appendix I: CeropegiaSPP* - Madagascan spp. only; Micranthocererrs(Brazilian endemic genus) plants in trade are likely tobe field-collected. M. auriazureus, the most heavilytraded species recorded in CITES statistics, is veryrare in the wild. M. dolichospermaticus is anotherspecies adversely impacted by trade.

Review of CITES listings for Succulents. Areview of the appropriateness of current CITES listingsfor succulent plants is required using the CITESappendices amendment criteria approved in 1994. Thisshould take into account information on conservationstatus in the wild, availability in trade, and extent ofcommercial propagation of both those plants currentlylisted on the Appendices and others threatened byinternational trade. The views of conservationorganisations, commercial growers, and botanic gardensshould be solicited.

Contact: IUCN/SSC Cactus and SucculentSpecialist Group

Astrophytum cactus nursery, Kurashiki, Japan.

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Pile of collected cacti, Terlingua, Texas.

4) Implementation of nursery registration. Allcountries with succulent plant nurseries should beencouraged to implement the resolution adopted at theNinth Meeting of the Conference of the Parties, Ft.Lauderdale, Florida, 1995, to CITES: Guidelines for theregistration of nurseries exporting artificially propagatedspecimens ofAppendix I species.

Contact: Appropriate CITES management authority

9 Regular review of CITES trade data forsucculents. As part of the ongoing Significant Planttrade process, regular analysis of the national reportedtrade in cacti and other succulents should be undertakenby the IUCN/SSC Cactus and Succulent Specialist Groupin association with WCMC and TRAFFIC International.Studies should determine the impact of trade onsucculents and their suitability for CITES listing.


6) Report on illegal trade in cacti and succulents.An annual report on international illegal trade in cactiand other succulents should be compiled and distributedto national authorities and relevant interest groups. Thiswill provide information on seizures and prosecutions andresults of investigations into availability of recentlydescribed and illegally obtained rare cacti.

Contact: TRAFFIC International

7) Training for CITES staff. It is important thatCITES Scientific and Management Authorities andcustoms agencies, in countries which have significanttrade in cacti and succulents, should be trained toimplement the trade controls and to recogniseappropriate illegal material and to deal with this trade.In-country training should be carried out in associationwith CITES field projects. Transfer training should alsobe encouraged, with personnel from exporting countriesseconded to CITES Authorities of importing countries.

Contact: CITES Secretariat

Ex situ con

8) The followingactions needing imconservation of succ

a>

b)

c)

d)

e)

f)

g)

h)

servatio

are priorities aplementationulents:

n

nd strategieswith regard

for future

Analyse existing ex situ collections and develop andenhance databases to track them. This can involvemore institutions and people than are presently servedin botanic garden databases, such as privatecollections and commercial nurseries. Developmentof such databases on a national or regional levelshould be considered a high priority as well as theimprovement of software documentation systems forcollections and protocols for the exchange databetween them.Develop a strategy for the use of ex situ collections ofcacti and succulents for conservation; that is, tohighlight priority species, then overlay these with c’xsitu collections enabling a clear identification ofparticular conservation needs.Develop means of exchanging data on cultivationtechniques and requirements to help promote CQ- situsurvival and propagation of rare and threatenedspecies.Develop educational programs at gardens for thegeneral public highlighting the need to conserve cactiand succulents.Conduct a study to determine how flooding themarket with artificially propagated plant materialwould affect the pressures on the wild populations.Develop a marketing strategy to sell artificiallypropagated rare succulents.Educate collectors about the conservation of wildpopulations and urge them to buy only artificiallypropagated plants.Develop more Species Recovery Programmes involvingmaterial available ex situ to act as models for otherconservation action, e.g. species reintroductionprogrammes.

Tour group, Desert Botanical Garden, Arizona.

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Increase awareness of CITES and its provisions forthe holders of cacti and succulent collections andpromote such collections as resources for theimplementation of this Convention. A CITESHandbook for Botanic Gardens has been published byBGCI (Akeroyd et al. 1994) and should be distributedwidely.Initiate research programmes to study the geneticvariability of ex situ collections of particularly rare andendangered taxa. Liaise with research institutions,universities, and NGOs.

Taxon specificAgavaceaeThe following are preliminary ideas. Furtherdevelopment of the p roposals i n co-opesuggested implementing agencies is needed.

ration with

9) Herbaria surveys of Agavaceae, sensu lato.Survey of herbaria holdings of all genera (with prioritygiven to Agave), to de te rmine the ex ten t o fdocumentation of the taxa. Specimens have been addedto various herbaria since the collections and herbariasearches of Gentry from the 1950s to the late 1970s.Surveys would reveal those taxa which are rare or are rarein the records, reveal distributional information and gapsin knowledge of a taxon’s distribution, and would help toclarify nomenclatural problems. A review of herbariaholdings may shed light on the status of missingsignificant specimens, especially type specimens.

Contact: DES, HNT and MEXU with Jardin Bot6nicode lnstituto de Biologia, Mexico City

10) Compile and provide a database of scientistsand herbaria actively researching groups withinAgavaceae. The initial steps in preparing this list (whichcould be linked with the review of herbaria holdings)would involve reviewing 1adex Herbariorum (Holmgren etal. 1990), consulting with the most recent edition of theInternational Register of Specialists and Current Research inPlant Systematics (Hunt Insti tute for BotanicalDocumentation), and consultation of recent editions ofthe IOS Repertorium, Bibliography Section. This databasewill assist in the development of field survey anddocumentation projects and subsequent management orrecovery plans for rare taxa.

11) Field survey and careful documentation of theAgavaceae, with Agave, Manfreda, Polianthes, andFurcraea as priorities. This is critical to the assessmentof the conservation status of any taxon. Following areview of herbaria holdings, it will become more clear asto which taxa and what areas need to be searched.Documentation of all Agavaceae, in particular Agave,Furcraea, and Yucca should be carefully undertaken and

Aloe ferox leaves which have been removed forextraction of aloe sap for commercial use in thepharmaceutical and cosmetic industries.

include photos, concise and descriptive notes (includinginflorescence size and shape), and floral measurements.Specimens should be processed in such a way as tominimise loss of character. Many specimens, with theexception of those prepared by Gentry and a few otherbotanists, are poorly prepared and provide littleinformation, thereby contributing little towards one’sunderstanding of the taxonomic group. In addition, sincemany collections of Agave occurred from the 1930s to theearly 1970s more recent collections, including areaswhere specimens were previously collected, should bemade. For example, Manfreda potosina is known fromthree collections, all of which were made between 1908and 1936. Its current status is unknown. Significant landutilisation activities or over-collection during the last SOor 60 years have probably impacted Agavaceaepopulations, and an assessment of their current status isrequired. Priorities for field survey are: Bahamas,Guatemala, certain states of Mexico - see regionalpriorities below.

Contact: Research instituand botanit gardens

tions such as u nivers ities

12) Identification of areas of highest diversity inAgavaceae, including protected areas, anddocumentation of the rare Agavaceae taxa foundwithin these areas. Action plans, which includeappropriate management and conservation strategies forthe rare taxa and their habitats should be developedwithin these identified areas.

Contact : IUCN/SSC Cactus and SucculentSpecialist Group

13) Review and conduct research on reproductiveprocesses and isolating mechanisms in Agave whichwill lead towards better understanding of thespeciation process in this group. A more flexibleapproach towards applying the general definition of a

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species rather than the more strict biological speciesapproach is required for such groups as Agave wherevegetative reproduction, hybridisation, and polyploidyplay very important roles in the speciation process. Suchan understanding may prevent problematic groups oftaxonomic and systematic interest (other than Fl hybrids)from “falling through the cracks” and thus not receivingrecognition or protection. Cultivation, selection andmigration of so-called species since remote antiquity needto be addressed in evaluating the integrity of species assome important fibre and pulque “species” may beremnants of selected and cultivated plants dating frompre-Conquest.

14) Development of an enc situ conservationprogram for Agavaceae. Botanic gardens provide anespecially effective setting for the study and conservationof the Agavaceae. Important ex situ collections of thefamily include those maintained by the Botanic Garden atUNAM, Mexico and the Desert Botanical Garden,Phoenix. A review of existing botanical garden holdings isof critical importance as recommended in Action 8a.Such a review should determine the number of specimensheld for a given taxon, whether they are from differentpopulations, and whether they are documented. Inaddition, a review of propagation schemes is essential.The ex situ conservation programme should emphasise theimportance of botanic garden collections for taxonomicand genetic research and the protection of crop geneticresources. Funding should be sought for ex situconservation activities in botanical gardens in a givenregion where rare Agavaceae occur, as part of a regionallyintegrated programme. For example, in Puebla, twobotanic gardens exist and can be involved in the ex situconservation of Agave peacockii, A. trianeularis, A. stricta,Beaucarnea gracilis, B. stricta, and Beschovneria calcicola.

Contact: Asociacibn Mexicana d e JardinesBothnicos

Aloaceae15) Development of a strategic conservation plan.In drawing up a conservation plan for the group, firstpriority should be given to the centres of diversity ofAloaceae, especially those in the southern and easternCape of Africa where many species are under threat. Aprerequisite for this project would be an analysis of thedistribution of aloaceous species in relation to the currentsouthern African protected areas. Education on the needfor habitat conservation and restrictions on collectingshould be an important component.


16) Medical research on Aloaceae. Surveys of theethnobotanical uses of alooid species should beconducted.

17) Fencing Aloe bowiea. This species is one of themost threatened of all the southern African A/W spp.Immediate conservation action, such as fencing off theremaining populations, is required. Alternatively, fivespecimens should be re-located to in situ and/or c-‘x situsafe sites.

Contact: Appropriate protected area managers

Crassulaceae18) Botanical exploration for this family in thefollowing regions: Central and South America, theNear East, southern Central Asia (Himalayas), EastAfrica, and Madagascar. Revisions of Madagascanspecies of Kalanchoe and Bryophyllrrm are urgentlyneeded.

Contact: Research institutions

Regional action proposalsCanary Islands19) Modification of species legislation. T h efollowing species should be added to Annex 1 of ‘Ordrensobre protection de especies de la flora vascular silvestrede la Comunidad Autonoma de Canarias’: Aeorziumsaundersii, Aichryson pachycaulon, Caralluma bwchardii,Euphorbia bourgeauana, and E. mell@-a.

20) Development of the proposed National Park onGran Canaria. The development of the proposedNational Park on Gran Canaria is urgently needed toprotect Euphorbia communities and eroded volcaniclandscapes which are threatened by land speculation andurbanisation.

Mediterranean region21) Review conservation status of theatened plantsin this region and evaluate all species with the newcategories of threat, in cooperation with the IUCNiSSCMediterranean Island Plant (see Delanoe et al. 19%) andEuropean Plant Specialist Groups.

22) Protect succulent habitat on the Atlantic coastalarea of Morocco near Agadir and to the south. Speciesinclude Euphorbia officinanrm, E. echinus, Car-allumaburchardii, C. europaea, and Kleinia a~ltel~~~lzordil~nl. Workmust be done to alleviate threats to these plants whichinclude industrial and tourism development, agriculture,and overgrazing.

Somalia23) Development of protected areas. It is importantthat when Somalia is in a position to develop a protectedarea system, sites rich in succulent plant species arcincorporated. These species are uniquely adapted to theclimatic and soil conditions and may be important inhabitat restoration. Although further botanical survey is apriority for parts of the country, including the Cal Madow

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hills of the north-east, there is sufficient informationavailable from earlier surveys on which to base siteselection. Ideally a Biosphere Reserve with strictlyprotected botanical sites within it should be createdwithin the north-east mountain region. Technicalassistance will be required and the generation of localsupport for conservation initiatives.

24) Taxonomic work in Asclepiadaceae.

Southern Africa(C. Hilton-Taylor)

25) Checklist of southern African succulents. Adefinitive checklist of southern African succulents needsto be compiled. This list should clearly define what ismeant by succulent and should include information onsynonymy, distribution, type of succulent (leaf, stem,caudiciform, etc.), plant use, conservation status, and keyliterature. Arnold and De Wet (1993) provide a startingpoint for such a list, but much of the information in thatvolume is inaccurate and out-of-date. An important stepwhich would greatly assist this and many of the otherprojects listed below, is the computerisation of allspecimen labels in southern Africa herbaria.

Contact: National Botanical Institute (NBI) andother herbaria in South Africa; National Herbaria inBotswana, Namibia, Swaziland, and Zimbabwe;systematists/taxonomists working on succulentgroups; the Succulent Society of South Africa; andthe Aloe, Cactus and Succulent Society ofZimbabwe

26) Production of a Red Data Book. A detailed RedData Book on southern African succulents needs to becompiled which has accurate information on populationnumbers and sizes and good documentation of the threatsto the species including quantitative estimates ofpopulation declines. An inventory of all taxa considered

s to be threatened in southern Africa was recentlypublished (Hilton-Taylor 1996b). Any new or correctedinformation should be sent to C. Hilton-Taylor at theaddress given in Annex 17.

Contact: NBI, provincial and national conservationagencies, professional and amateur botanists

27) Review of in situ succulent plant protection. Thereis a need to determine which species and the number oftheir populations are effectively protected in the existingprotected area network. Distribution data for all thesucculent taxa should be analysed using iterativeprocedures as’ described by Rebel0 (1994) to obtain theoptimal reserve configuration for preserving themaximum diversity of southern African succulents. Thisconfiguration needs to be compared to the existingprotected area network and gaps in the networkidentified. All the high priority areas, particularly major

sites of succulent diversity, need to be declaredconservation areas. A number of gaps in the protectedarea network have already been identified (see chaptersin Huntley 1989 and 1994). Hilton-Taylor and Le Roux(1989) have indicated a number of areas in the SucculentKaroo which need to be set aside for the protection ofsucculents, however, few of these recommendations havebeen acted on.

Contact: Conservation agencies, NGOs, herbaria

28) In situ succulent plant conservation inZimbabwe. Kimberley (1991) has outlined an ambitiousplan to select thirty habitats in Zimbabwe for botanicalreserve purposes. The findings of Timberlake and Muller(1994) should be incorporated into this plan. Kimberley(1991) estimated that the plan would cost approximatelyfive million Zimbabwean dollars (equivalent at that timeto one million U.S. dollars) extended over a ten yearperiod. This plan would also have the added economicbenefits of creating employment for at least sixty, if notmore, people. The proposed plan for Zimbabwe shouldbe emulated in other southern African countries.

Contact: The NBI and other southern Africanbotanical institutions are in the best position to carryout the analysis, but the purchase and setting asideof land is the responsibility of national governmentsand government conservation agencies with the aidand support of NGOs such as the SucculentSociety, Wildlife Society, and World Wide Fund forNature.

29) Participation in protected area planning. InSouth Africa during the proposed redistribution of land tothose people with historical claims on certain areas it willbe important to demonstrate the tangible benefits ofconservation to the community concerned througheducation, workshops, etc., especially aspects such aseconomic benefits and job opportunities, so as to ensurefull public participation. Archer (1993) for example,describes the participation process followed innegotiations between pastoralists and conservationists inthe recently established Richtersveld National Park, hometo many endemic species of succulent plants.

30) Conservation on private land. Alternatives topublicly owned or legally designated conservation areasare urgently required in situations where formal protectedarea status is inappropriate. Farmers should be madeaware of the importance of conservation and they shouldbe encouraged to leave parcels of natural land for theprotection of as wide as possible a variety of geneticdiversity. In South Africa, landowners may on a voluntarybasis register part of their land as a Natural Heritage Sitewith the Department of Environment Affairs, so as toprotect important natural sites (Fuggle and Rabie 1992).The landowner may receive some management advice forthe site, but retains full rights over the property, and asthe registration has no legal status, it falls away when the

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landowner dies or when the property is sold. Anotheralternative is for a landowner to join a conservancy wherethe owners of several properties have combined resourcesfor the improved conservation of the natural areasremaining on their land (Fuggle and Rabie 1992). These

be introduced. There are many examples of such‘stewardship’ schemes world-wide.

Contact: Government departments, conservationagencies, NGOs, landowners, and publicparticipation

31) Survey of ex situ succulent plant conservation.Which species, how many, and the sources of all thematerial, are in cultivation in botanical gardens insouthern Africa and elsewhere in the world needs to bedocumented. In addition to botanical gardens, the speciesin large privately owned collections particularly of certainsucculent genera, also need to be recorded. All botanicalgardens and individuals with important collections shouldbe encouraged to register as holders of an 10s GenericReserve Collection.

Contact: Botanic Gardens ConservationInternational (BGCI), NBI, all botanical gardens insouthern Africa, Succulent Society of South Africa,National Cactus and Succulent Society ofZimbabwe, IOS

32) Improving CITES implementation. In terms ofregulating international trade in succulent species fromsouthern Africa, it may be necessary to propose the listingof additional taxa on the CITES Appendices. This will bedetermined from the results of trade surveys conducted byTRAFFIC. Such surveys will have to be conducted atfairly regular intervals to monitor trends. A number of

additional aspects concerning CITES also need to beattended to: (a) non-member states within southernAfrica should be encouraged to join; (b) CITESregulations need to be correctly and efficiently enforcedby each state and especially by the different provinceswithin South Africa; (c) inspectors need to be givenadequate training and literature to help them in theidentification of CITES listed material; (d) there shouldbe separate Management and Scientific CITESAuthorities for each country; and (e) the fate ofconfiscated material should be clearly defined in a policydocument.

Contact: CITES Secretariat, TRAFFIC, governmentconservation agencies, botanical research institutes

33) Development of national legislation. Thedeficiencies in national and provincial conservationlegislation as outlined in the Southern African RegionalAccount need to be addressed as a matter of top priority.For example, (a) Botswana urgently needs legislation toprotect its flora; (b) government funding for the PPC inLesotho is totally inadequate and staffing is insufficient;(c) the list of protected plants in Namibia is in need ofrevision (M. Strobach, pers. comm.); (d) now that SouthAfrica is divided into nine provinces, each will have todraw up its own conservation legislation, providing a goodopportunity to update and tighten up the ordinances andto ensure even standards and treatment across thecountry; and (e) penalties imposed for offences are inmost cases far too low to act as a deterrent and should besubstantially increased to match those for offencesrelating to animals.

Contact: National governments, conservationagencies, botanical research institutes,environmental lawyers

A koker boom (Aloedichotoma) forestwhich could easilybecome a major touristattraction.

34) Law enforcement. The provision of appropriateconservation legislation must also ensure adequate lawenforcement. South African legislation for the protectionof plants in general and succulent plant species inparticular is generally satisfactory, but in many cases theenforcement falls far short of what is desirable. This isnot a reflection of the actual abilities of the law enforcers,but on their small numbers and the vast areas they have topolice (Cowling and Olivier 1992). The main reason forthis, according to Kimberley (1991) is the lack ofpersonnel, and especially the absence of any active andfull-time inspectorate, which is largely attributed to theshortage of sufficient financial support.

Contact: National governments, conservationagencies, honorary conservation officers

35) National Biodiversity Action Plans.Government funding for flora conservation in everysouthern African country is inadequate. This issue shouldbe addressed by countries when compiling national plansin accordance with the Convention on BiologicalDiversity. Namibia is probably the first southern Africancountry to have started this process (G. Maggs, pers.comm.).

Contact: National governments, conservationagencies, botanical research institutes

36) Education and public awareness. As most landin southern Africa is privately or communally owned anextensive and intensive environmental education andawareness campaign using all the media is required toeducate the general public about our floral wealth andespecially those species at risk (see Smith 1994). Thecampaign needs to instil in the population of eachcountry, particularly people in the rural areas anddevelopers, a corporate responsibility to protect andsafeguard its floral heritage for future generations. TheCommunal Areas Management Programme forIndigenous Resources (CAMPFIRE) in Zimbabwe couldserve as a role model for such an education campaign.The CAMPFIRE project was started by the Zimbabweanconservation authorities in 1982 as a result of theincreasing conflict between people and wildlife and therealisation that much of the land was better used forconservation than for conventional economically andecologically unsound agricultural purposes (Holt-Biddle1994). Similar pilot schemes have been started inBotswana and Namibia, but they have not had the samemeasure of success as CAMPFIRE (Holt-Biddle 1994).

Contact: National governments, conservationagencies, education authorities, traditional leaders,universities and other educational institutions,NGOs

Awareness of the floral wealth also needs to be extendedto those people in authority, especially those directlyresponsible for the conservation of the flora. Ever since

the concept of conservation gained popularity andmomentum in southern Africa, the emphasis has largelybeen on the fauna (Huntley 1978; Kimberley 1991). Thelack of care and expertise about the flora has been suchthat those in authority often do not know one protected,specially protected, or endangered plant species fromanother.

Contact: National governments conservationagencies, the NBI and other botanical institutes,educational institutions, NGOs like the succulentsocieties, the media

37) Promotion of the financial value of succulents.There is a growing realisation in southern Africa thatsustainable and imaginative use of natural resources canbe used to generate considerable income, especiallythrough the concept of ecotourism (Cowling 1993).Unique areas of succulent diversity could be used togenerate income, especially if promoted as touristattractions. Millions of visitors to Arizona in the USA forexample, pay to see the representative succulent plantcommunities (organ pipe cactus, giant Saguaro cactus,and Joshua trees) which are protected in the SonoranDesert. The kokerboom (Aloe dichotoma) forests of theNorthern Cape and Namibia, Euphorbia thickets of theEastern Cape, Sesamothamnus lugardii populations inBotswana, and Lithops colonies in Namibia are primeexamples of such resources which could easily bepromoted.

Contact: Conservation agencies, NGOs, localauthorities, and tourism organisations such asSATOUR

38) Development of succulent plant nurseries. Tohelp prevent the removal of succulent plants from thewild, attempts are required to establish sufficientnurseries to meet the demand for plants. Such nurserieswould not only supply plants and seeds for thehorticultural trade but also plants used for traditionalmedicinal purposes. Such nurseries should operateaccording to the guidelines set out in the 10s Code ofConduct (Oldfield 1990). Rural communities inparticular, should be helped and encouraged to set upsuch nurseries, e.g. in the Richtersveld or in Lesotho forAloe polyphylla (see Talukdar 1983). These nurserieswould not only benefit the local community financially butwould hopefully educate people about the value of theplants thereby engendering a conservation ethic.

Contact: Conservation agencies, NGOs, traditionalleaders and their rural communities, commercialnurseries, traditional healers, TRAFFIC, the NBI andother botanical institutes

39) In situ monitoring of threatened species. Long-term monitoring studies in situ of selected threatenedsucculent species and their habitats are required to gainfurther insights into the conservation requirements of

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endangered species. Such studies could be modelledalong the lines of those conducted on cacti in Mexico byCan Te, A.C. in collaboration with CITES. Haworthiaand Euphorbia are two prime genera which have manysuitable candidates for such a monitoring programme.

Contact: CITES, conservation agencies, SucculentSociety of South Africa, National Cactus andSucculent Society of Zimbabwe, the NBI, universitybotany departments

40) Rescue operations for threatened succulents.As part of the ex situ conservation measures, a co-ordinated approach is required across southern Africa forthe rescue of succulent plants, especially threatenedspecies, from areas intended for development. Theserescued plants should be grown in cultivation with theview to reintroduction to another suitable and safe site ormoved immediately. The Botanical Society of SouthAfrica has a ‘Search and Rescue’ group run under theauspices of their Flora Conservation Committee. Thisgroup, however, is purely Western Cape based and dealsmainly with in situ problems of habitat conservation,although some plants have been removed from the wildand taken to Kirstenbosch National Botanical Garden forcultivation. ‘Operation Wildflower’ is another suchgroup, which in recent years has only operated in theTransvaal. This group focuses mainly on indigenoussucculents (aloes) and bulbs, which are removed fromareas slated for development and cultivated in privategardens. While these local efforts are praiseworthy,national or possibly regional, well co-ordinatedapproaches to the problem are required, especially ifeffective reintroduction programmes are to be started.The Centers of Plant Conservation in the USA couldpossibly serve as a role model for such a programme (Falk1992).

Contact: Conservation agencies, botanical gardens(especially the NBI), Botanical Society of SouthAfrica, Succulent Society of South Africa, Cactusand Succulent Society of Zimbabwe, other NGOs

Madagascar41) Incorporation of succulent plant expertise in theIUCN/SSC Specialist Group for Madagascan plants.There is an urgent need for conservation activitiesrelating to the plants of Madagascar to be moreeffectively co-ordinated through the creation of aspecialist group of experts from Madagascar andelsewhere, which includes expertise on the succulent flora.

Contact: SSC Plants Programme

42) Analysis of the distribution of succulentspecies in relation to the current and proposedprotected areas. GIS mapping of the knowndistribution of succulent species by genera should becompleted, based on the botanical literature and theresults of the CITES field project, to highlight the centres

of diversity for each succulent genus. Compare centres ofdiversity for each genus and the protected area network.Recommend for expansion the protected area coverage ofthe main centres of diversity for succulent plant species.A GIS project based on orchid, palm and legume data isalready underway under the auspices of the Royal BotanicGardens, Kew. A technician has been employed in Paristo enter locality data for this project. Support should beprovided for similar data to be accessed for the succulentflora.

Contact: The Royal Botanic Gardens, Kew

43) Detailed assessment of the conservation needsof the xerophytic vegetation of south-westMadagascar. A comprehensive survey should beconducted of the spiny thicket vegetation of south-westMadagascar to determine the best remaining sites forconservation purposes. Site selection can be based onrepresentative examples of the vegetation structurallydefined, succulent species richness, and presence of rareand/or threatened endemics.

Contact: WWF-Madagascar/ ZSS

44) Cap Sainte Marie Special Reserve. This reserveis important for the protection of the endemic andthreatened Euphorbia cap-saintenzaI’it’l%siS and for othersucculents such as Aloe millotii, Alluaudia comos~~, andAlluaudiopsis fi’herensis. While some information on thespecies present is available, a full botanical inventoryshould be carried out. The reserve is also scenicallvattractive and, although no visitor resources exist atpresent, it is potentially an important site for ecotourism.Reassessment of the boundaries, exclusion of seriouslydegraded and botanically less important areas and fencingto exclude livestock from the core area is necessary. Thedevelopment of ecotourism and area management plansshould be encouraged.

45) Development of a ‘lItlear botanical garden’.Development of a botanic garden with natural vegetationprotected as a reserve in the vicinity of Tulear.

Contact: University of Tulear

46) Assessment of Strict Nature Reserves.Assessment of the condition and realistic prospects formedium- to long-term protection of succulent plants atReserve naturelle integrale de Tsimanampetsotsa andReserve naturelle integrale d’Andohahela (parcel 2).

Contact: Missouri Botanic Garden

47) Detailed survey of the western escarpment ofthe Central Plateau. The area between the Mangokyand Manambaho Rivers is in need of a detailed survey toassess the conservation status and conservation needs ofthe succulent flora.

Contact: WWF-MadagascarlZSS

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Echinocereus chisoensis, Big Bend National Park,Texas. Vulnerable.

48) Creation of new protected areas. Protection ofthe priority sites for succulent plant conservationidentified above is urgently required. A feasibility studyshould be carried out to determine the best form ofprotection, site boundaries, management requirements,willingness of local people to support protected areas andthe castings involved. Funding proposals should bedeveloped for the creation of reserves where theagreement in principle of all interested parties has beenreached.

49) Development of plant conservation legislation.

Contact: Government offices, lobbying groups

50) Improvements in nursery monitoring. It isimportant to have closer monitoring of plant collectionand production for export in the commercial nurseries.There are currently around six such establishments.Details of stock plants and their origin, levels ofproduction, and trade by species should be maintained.Procedures for the monitoring of nurseries by regularinspection and review of documentation should beestablished. Training for a member of staff withresponsibility for control of plant exports should bearranged.

Contact: Department des Eaux et Forets (DEF)

51) Development of a CITES Scientific Authorityfor plants in Madagascar.

52) Popular guide to the succulent plant species ofMadagascar for local use. This will promote localinterest in conservation in plants and can be used inschool education.

53) EX situ conservation at Part botanique etzoologique de Tsimbazaza. The park is now equippedwith nursery facilities suitable for propagation on a largescale. The garden needs to produce a strategy for the ex

situ conservation of succulent plants. Cultivation of rarespecies should concentrate on high altitude plants fromcentral Madagascar and garden staff should beencouraged to look at the natural localities of the plantsin cultivation to ensure that horticultural practices areappropriate. There has been a renovation programmefocused on the rocker-y which displays the succulent floraof Madagascar. Construction is almost complete; plantingwill require further work for several more years. There isan urgent need for explanatory signs and leaflets on thegenera, their natural habitats, and cultivation notes; andtraining in the ecological requirements of native species,

E practical conservation, and display of succulents.

a> 54) Construction of a database of the existing$ succulent collection at Toliara. An importantz collection of natural source succulents is maintained at

the privately owned Arboretum d’Antsoky. The scientificvalue of this collection should be evaluated and, ifappropriate, funding provided for a technician to developa database on site. This could be linked to Action 8a.

India55) Survey of endangered species. Survey andinventory of endangered succulent species as a basis forselection of reserves. Tea companies, owners of vasttracts of land rich in succulents, could also be involved asthey are in a position to set aside small areas as reserves.

Contact: President of the Cactus and SucculentSociety of India

56) Sustainable use management plan. Manyspecies are collected for local and commercial use and arethreatened by over-collection. Attention should be givento educating users about sustainable collection methods.

United States of America57) Implementation of published recovery plans.The US Fish and Wildlife Service (USFWS) oversees andpublishes status reports of recovery plans. See Chapter 2- National Legislation for details. Although the onus is onthe USFWS, outside individuals and institutions can helpwith implementation.

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Reading an electronic chip.

58) Peyote salvage operations in southern Texas.Peyote populations in south Texas are of considerableimportance to members of the Native American Church,who obtain plants to use as their sacrament. Most of theland is privately owned, so often neither NativeAmericans or Peyoteros (those who have permits tocollect peyote) are allowed onto the land where the plantsgrow. Consequently available populations are being badlyovercollected. Another serious threat to the peyote insouth Texas is the root plough which is used by landowners to remove the natural plants before sowing exoticplants for grazing. If land owners could be persuaded topermit peyoteros to collect the peyote plants prior toploughing, these cacti could be transplanted to anothersite to be grown for traditional use. The development ofnursery seed grown peyote stock for local use should alsobe investigated.

Budget: Funding would provide for the lease ofland for growing the cacti and to hire peyoteros tocollect them. A pilot project should be carried outand funded for $5000 - 10,000.

59) Long-term monitoring of Echinocereuschisoensis. This taxon is considered threatened in theUSA. It grows only in Big Bend National Park, Texas, butthere may be additional populations outside the parkboundaries in Texas and similar plants are known inMexico. Survey is needed to determine the range of thisspecies. Permanent monitoring sites should also beestablished to study the long-term dynamics of the knownpopulations, especially those protected within the Park.

Budget: Annual cost would be $2000 - 3000.

60) Establishment of permanent monitoring sitesfor species of Sclerocactus and Pediocactus in theUSA. These genera contain some of the rarest cacti inthe USA. Permanent monitoring sites should beestablished to study the population dynamics of theseplants, as well as to determine the effects of activities suchas collecting, mining and construction. Some of this work

should befor several

done in cospecies are

-0perati on with Native Americalocated on Indian reservations.

ns,

Budget: The estimated cost of such work is $3000 -5000 per year.

61) Search for Astrophytum asterias populations.This species is known from a single locality in Texas, butalmost certainly it was once widespread throughout theshrubland of south Texas and into north-eastern Mexico.It is known to occur east of the Sierra Madre Oriental andsouth of Ciudad Victoria, Tamaulipas in Mexico and islikely to occur elsewhere. Much of the land where itgrows, or has grown, is being developed for agriculture. Asurvey of distribution and conservation status is required.

Budget: The estimated annual cost would be$4000.

62) Support for succulent conservation work byinstitutions working with the Center for PlantConservation (CPC). Various cacti and othersucculents are currently being propagated and studied bymember institutions of CPC; however, their funding islimited, and financial support for their important fieldwork, as well as conservation of germplasm, is muchneeded. More publicity is needed about CPC’s programand how botanic gardens and specialists can participate inthis.

63) Assistance in the development of conservationeducation. The public must be informed of the presentthreats to many plant species in the wild. Some USinstitutions have plans to develop extensive programmes.For example, the Desert Botanical Garden has developedplans for a Conservation Trail by which the public will beinformed of the purpose of conservation and how it mavbe accomplished. Outside funding to encourage suchprogrammes is required.

64) Survey of Agavaceae. This work is needed inArizona and Texas.

Contact: Desert Botanical Garden

Epithelantha micromeris, Cuesta La Muralla, Mexico.

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Mexico and United States of America65) Cooperative Mexican-American conservationstudies along the US-Mexico border. Much of thearea south of the Rio Grande, especially that area to thesouth of west Texas is poorly known. Several rare cactimay occur in this area but extensive field work isnecessary to determine ranges of some of these plants,such as peyote (Lophophora williamsii), Ariocarpusfissuratus, Echinocereus chisoensis, and Sclerocactus(Neolloydia) mariposensis.

66) A survey of the genus Ferocactus in trade. Astudy should be conducted to compile data from the rangestates in Mexico and the USA and the major importingcountries of Europe and Japan. A sizeable trade exists inseed-grown Ferocactus and the study must distinguishbetween wild and nursery-grown plants.

67) Epithelantha micromeris. A population survey ofthis species is needed to determine the status of thisspecies in the wild in Texas and Mexico. It can be growneasily and seedraised plants are generally seen inEuropean nurseries. Wild-collected plants are, however,also available in trade and the impact of internationaltrade on wild populations needs to be assessed.

68) Permanent marking of rare plants. Illegalcollecting continues to be a significant threat to somespecies of cacti in Mexico and the USA. It would behelpful to mark individual plants in some of theseseriously threatened populations, not only to enableauthorities to determine the origin of confiscated plants,but also to assist researchers in locating the plants in thefield. Some of the most popular plants are small andnearly invisible, and permanent markers would greatlyfacilitate continuing studies of these populations.Electronic chips could be inserted into the body of thecactus without injury to the plant.

Budget: Equipment and supplies would cost $5000- 6000.

Mexico69) Education. Long term conservation can be servedbest through education at the grass-roots level. Mexiconeeds books about cacti and other succulents, suitable foruse in basic education. A series of children’s books onappreciation of arid habitats and the native flora is now inpreparation by Can Te, A.C. Other planned publicationsinclude a pictorial encyclopedia of Mexican succulents,introductory and primarily pictorial works on succulent-rich regions, and field identification guides to potentiallythreatened taxa. Continued support is needed in this veryimportant area. While it will remain difficult to controlthe collection of succulent plants by individuals fordecorative or medicinal purposes, educational effortsshould help.

70) Preparation of c lear guidel ines onconservation legislation and collecting regulationsrelating to Mexican succulents. A major problem inMexico is a lack of communication about conservationactivities and lack of understanding about the federal lawsthat affect native plants, especially important among thefarmers and villagers who live in the arid regions ofMexico. Local people need to be informed ofconservation activities, especially when they involvepopulations of plants occurring near their farms orvillages, especially if the succulents are found on theirejidos (communal land tenure system). The success oflocal conservation efforts is possible only with theknowledge and support of local residents.

71) Enforcement of conservation legislation.Mexico has called on importing countries to help withproblems of illegal plant exports. Recently, Mexicanauthorities have been studying where collectors go withinthe country and concentrating enforcement efforts inthose areas. Complementary stricter controls byimporting countries are required.

72) Indigenous plant propagation. The MexicanGovernment is encouraging those botanical gardens andnurseries that are beginning to propagate succulents forboth domestic and international markets. In this way asource of income is generated, and the pressures on wildpopulations of succulents are reduced. If inexpensiveplants of high quality can be marketed by responsibleMexican nurseries, this should lead to less demand onfield-collected (and illegal) cacti and succulents.Propagation activities and market research clearly meritcontinued encouragement and support from both theMexican authorities and international conservationorganisations. Assistance with mass-propagationtechniques and marketing skills is required. Internationalexpertise in the development of the horticultural industryshould be offered.

73) Development of protected area system. Prioritysites for succulent plant conservation have been identifiedin the Regional Account for Mexico. These sites will bebrought to the attention of the relevant authorities inMexico and investigations carried out to identify the mostappropriate means to protect the sites.

74) Population studies. Ecological and populationstudies are the foundation of future conservationactivities. All recommendations for conservation actiondepend, at the least, upon reliable population data inorder to best allocate limited funding. The followingproposed population studies have been proposed for theAction Plan by Can Te, A.C. Only plants that are felt towarrant study as candidates for Critically Endangered,Endangered, or Vulnerable status (new IUCN threatcategories) are included in the examples. Each studywould provide a population estimate of sufficientaccuracy for the plants listed to allow threat classification

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by SEMARNAP. Each study corresponds to a one weekundertaking for three investigators. The cost for eachstudy (US $740) covers per diem expenses andtransportation costs.

Las Tablas - Cuidad Maiz - Los Cerritos, State of SanLuis Potosi

Coryphantha maiz-tablasensisMammillaria aureilanata (1 of several sites)M. microtheleM. dumetorum (1 of several sites)Turbinicarpus gielsdoeianusT. knuthianusT. lauiT. lophophoroides

Zaragoa, State of Nuevo Le6nEcheveria shavianaEcheveria sp. nov.Echinocereus knippelianus var. reyesiiMammillaria rubrogandisM. glassii var. novaThelocactus conothele (yellow-flowered form)Turbinicarpus subterraneus var. zaragosae

Aramberri, State of Nuevo Le6nAriocarpus kotschoubeyanus (white-flowered form)Mammillaria albicoma (1 of several sites)Pelecyphora strobiliformisThelocactus conothele var. aurantiacusTurbinicarpus hoferiT. pseudopectinatus (red-flowered form)T. subterraneus var. subterraneusT. schmiedickeanus var. dickisoniaeT. schmiedickeanus var. gracilis

Galeana, State of Nuevo Le6nAztekium hintoniiGeohintonia mexicanaMammillaria glassii (1 of several sites)M. weingartiana (1 of several sites)Turbinicavus pseudopectinatus (red-flowered form)T. booleanus (aff. T. subterraneus)

Ascension, State of Nuevo Le6nEscobaria roseana var. galeanensis (1 of several

sites)E. asperispina (2 of several sites)Echinocereus knippelianus var. krugeriE. pulchellus var. sharpiiMammillaria glassii var. ascensionisM. weingartiana (1 of several sites)Thelocactus conothele var. argenteusTurbinicarpus sp. nov. (aff. T. gautii)

Rayones, State of Nuevo Le6nEscobaria roseana var. galeanensis (1 of several

sites)

Agave victoriae-reginae (1 of several sites)A riocarpus staph iros trisAztekium ritteriEcheveria lilacinaEchinocereus knippelianus (new form)E. pulchellus (new form)Turbinicarpus swobodae

These examples cover abbreviated field investigations.feasible as a consequence of prior experiences of the Cm

Te, A.C. staff. Similarly abbreviated field investigationsshould be possible as a consequence of prior cxpericnccof the UNAM staff in the Tehuactin area and theUniversidad Aut6noma de Tamulipas staff in theJuamave area. Additional proposals can be solicited fromother institutions in Mexico, either directly or throughCan Te, A.C., in order to take advantage of both work inprocess and proximity to the sites.

With the participation of all relevant institutions incarrying out population estimates of CriticallyEndangered, Endangered, or Vulnerable succulents, itappears that the task can be completed within two years.Plant populations are dynamic, and continuingobservations will be necessary to evaluate new potentiallythreatening situations.

75) Surveys of Agavaceae. The following arc prioritvisurvey projects for Agavaceae. Organisations serving aspossible primary contacts are suggested, thcsc indicatedby their herbarium acronyms and appropriate botanicalgardens. These suggestions do not constitute acommitment by that institution at this time.

a>

b)

4

4

Field surveys and documentations of Aglr~ in Baja.California (including islands)

Contact: DES and Desert Botanical Garden withBCMEX and El Charco del Ingenio, San Miguel deAllende, GuanajuatoBudget: $15,000 for 5year period

Field surveys and documentation of rare Agavaceac inSonora

Contact: DES and Desert Botanical Garden with ElCharco del lngenio and Centro Ecologia de Sonora.

Field surveys and documentation of rare Agavaceac inCoahuila

Contact: DES and Desert Botanical Garden, with ElCharco del lngenio

Field surveys and documentation of Aga~ and rareAgavaceae in Oaxaca

Contact: MEXU and Jardin Bottinico de lnstituto deBiologia with OAX, and El Charco del lngenio

139

e) Field surveys and documentation of rare Agavaceae inSan Luis Potosi

Contact: SLPM and El Charco del lngenio

f) Field surveys and documentation of rare Agavaceae inJalisco

Contact: MEXU and Jardin Bot6nico de lnstituto deBiologia with IBUG, and El Charco del lngenio

Other states of lower priority, but nonethelessrequiring field surveys and documentation for rareAgavaceae, are listed below with suggested contactorganisations along with MEXU and Jardin Botanic0 deInstituto de Biologia and El Charco de1 Ingenio:

a) Durango, with CIIDIR and DES and DesertBotanical Garden

b) Nayaritc) Hidalgod) Vera Cruze) Mexicof) Pueblag) Guerrerh) Chiapas, CHIP

Central America76) Field surveys of Agavaceae in Guatemala.

Contact: AGUAT and UAT (Beaucarnea)

Caribbean77) Survey of the conservation status of the generaAgave and Furcraea in the Bahamas. Theconservation status of the endemic species ofAgave in theBahamas is currently unknown. Review of the status ofwild populations and current threats is required in orderto effectively implement the SPAW Protocol of theCartagena Convention.

Flower of Leptocereus wrightii, a species practicallyextinct in the wild.

Cactus scrub habitat near Azu&, DominicanRepublic.

78) Creation of an SSC Specialist Group for theFlora of the Caribbean islands. There is an urgentneed for conservation activities relating to the plants ofthe Caribbean Archipelago; not only are species of cactusand other succulents facing extinction, but also many taxawithin the Arecaceae, Myrtaceae, Rubiaceae,Melastomataceae, Magnoliaceae, Eriocaulaceae,Xyridaceae, and other families. In a multinational regionsuch as the West Indies, the best way to makeconservation activities effective is by co-ordinating themthrough the creation of an international Specialist Group.

79) Detailed assessment of the conservation statusof cacti and succulent species that remaininsufficiently known, based on field survey. Prioritiesare:

80) Detailed search for the ‘extinct’ or nearlycd extinct cactus and succulent species. A detailed=i search for the most endangered taxa in the wild is3 necessary to determine whether they are gone or are yetg capable of surviving. Mapping of the last remaining sites5 of these taxa, if existing, and assessment of their’ conservation needs is urgently needed. Taxa to be

considered:

.*

140

a) Agave brevipetalab) A. brevispinac) Neobuchia paulinaed) Opuntia ekmaniie) 0. falcataf) Dorstenia caimitensisg) D. cordifoliah) D. crenulatai) D. erythrandraj) D. flagelliferak) D. hotteana1) D. marginatam) D. multisquamean) D. tuberosao) Cissus haitiensia

Contact: NYBG, DNP, JBN-RMM

a)b)44e>f >g>h)9.J>90I-4

Escobaria cubensisHylocereus cubensisLeptocereus scopulophilusL. weigh tiiMelocactus actinacanthusM. matanzanusOpuntia borinquensis0. corallicolaPereskia quisqueyanaSelenicereus innesiiBorrichia cubanaCnidoscolus fiagransC. quinqueloba tus

Contact: NYBG, IES, MNHN, JBN, DNP, DRN

81) Analysis of the distribution of succulentspecies in relation to the current and proposedprotected areas. Mapping of the known distribution ofcacti and succulent species based on botanical literatureand field work will highlight the centres of diversity. Acomparison between these centres of diversity and theprotected area network is to be made. Also,recommendations should be made for the expansion ofprotected area coverage to include all the centres ofdiversity for cacti and succulent plant species, and most, ifnot all, of the taxa involved.

Contact: NYBG

82) Detailed assessment of the conservation needsof important dry shrubwoods and semi-desertcactus communities not covered by nationalprotection systems. A comprehensive survey ofimportant unprotected dry shrubwoods and semi-desertcactus communities is needed to determine the bestremaining sites for conservation purposes. Site selectionfor the expansion of protected area coverage should bebased on representative examples of the vegetationdefined by structure, succulent species richness, and thepresence of rare and/or threatened endemics. Prioritiesare:

a)

b)

c)d)

e)

f)

g)

h)i)

Coast and lowlands from Guantanamo to Maisi,CubaCoastal terrace between Gibara and Puerto Padre,CubaHolguin serpentine outcrops, CubaLake Enriquillo Valley, Domincan Republic, Cul-de-Sac Plain, HaitiShrublands between Bani and Azua, DominicanRepublicCoast and lowlands from Mule St. Nicolas to Port-de-Paix, HajitiCoast and lowlands between Treasure Beach andLittle Pedro Point, JamaicaHellshire Hills coastal and lowland area, JamaicaDry coastal plains in the Turks and Caicos Islands,Bahama Archipelago

j) Eleuthera Is land rocky p la ins , BahamaArchipelago

k) Great Inagua sandy and rocky flats, BahamaArchipelago

1) Long Island shrublands, Bahama Archipelago

Contact: NYBG, COMARNA, IES, DNP, NRCD

83) Creation of new protected areas. Protection ofall the priority sites for cactus and other succulent plantsidentified in the Action Plan is urgently required. Afeasibility study should be carried out to determine thebest form of protection, site boundaries, managementrequirements, requirements of local people and the costsinvolved. Most of the sites proposed are small in terms ofarea coverage (for example, Cerro de San Francisco, inthe Dominican Republic, is only a hill). Thus, theirselection follows the general strategy conceived under“Priority sites for conservation”. Funding proposalsshould be developed for the creation of reserves wherethe agreement in principle of all interested parties hasbeen reached. Proposed sites are:

a>b)4

d)e>f)g>h)

9.J>

k)9

m)n>4

Santa Cruz de1 Sur lowlands, CubaSierra de Anafe, CubaSierra de Somorrostro and neighbouring hills,CubaSierra de Najasa, CubaTres Ceibas serpentine outcrop, CubaJatibonico ophiolitic outcrop, CubaBayahibe coast and lowlands, Dominican RepublicCerro de San Francisco, Banica, DominicanRepublicTetas de Cayey mountain ridge, Puerto RicoNorthern Grand Terre of Guadeloupe, LesserAntillesBarbuda lowlands, Lesser AntillesSouth-eastern peninsula of St. Kitts, LesserAntillesLa Soufriere, St. Vincent, Lesser AntillesThe Bluff, Cayman Brat, Cayman IslandsEast End, Little Cayman, Cayman Islands

Contact: NYBG, MINAGRI, COMARNA, DNP, DRN,NRCD, NTCI

84) Assessment of strict nature reserves. M a n yprotected areas are not adequately protected on theground. Assessment of the condition and realisticprospects for long-term protection of cacti and othersucculent plants in the existing protected areas of theCaribbean Islands is to be done. Recommendations tosecure the protected status of the plants involved shouldbe given.

Contact: NYBG

85) Detailed survey of the Department duNord’Ouest, Haiti. In all the Caribbean the dryshrublands and coastal areas of north-west Hai’ti are the

141

poorest known sites with respect to their native succulentflora. Judging from old collections of this area, thereseems to be an undescribed new genus of Cactaceae. Theregion is in great need of a detailed survey to inventorythe succulent taxa, and to assess the conservation statusand conservation needs of its succulent flora.

Contact: NYBG, Fundation botanique d’Ha’iti

86) Development of plant conservation legislation.Each Caribbean country should develop plantconservation legislation to give special protection to itsrare and threatened species.

87) Development of CITES Scientific Authoritiesfor plants with a mechanism for regionalcoordination.

88) Completion and amendments to the succulenttaxa of the Caribbean’s SPAW Protocol Annexes.

89) Improvements in nursery monitoring. Closermonitoring of plant collections and production for exportin the commercial West Indian nurseries is important.Details of stock plants and their origin, and levels ofproduction and trade by species should be maintained.Procedures for the monitoring of nurseries by regularinspection and review of documentation should beestablished. Training within the CITES ManagementAuthority for a member of staff with responsibility forcontrol of plant exports should be arranged.

90) Ex situ conservation. Creation of a complete exsitu Caribbean succulent plant conservation collection isurgently required. Due to the potential for adversenatural environmental factors in the West Indies, such ashurricanes, it is desirable to have more than one regionalcollection. The St. George Village Botanical Garden onSt. Croix (US Virgin Islands) has, at present, the mostimportant ex situ collection of endangered Caribbeancacti. Also the Jardin Botanic0 National at Havana, andthe Cayman National Botanic Park in Grand Caymanhave started, on their own initiative, ex situ succulent plantconservation collections.

The idea of creating a sanctuary for endangeredCaribbean succulents at the Tetas de Cayey mountainridge (Puerto Rico) by its private owner is worthy ofbacking and support. The development of such asanctuary in a place with natural vegetation protected as areserve (the rocky slopes are inhabited by a yetundescribed variety of Melocactus and several localendemics), is a very exciting idea. There is an ongoingproject for the rockery displays. This place is soon goingto be equipped with nursery facilities suitable forpropagation on a regular scale.

91) Educational programs. There is a need todevelop educational programmes for the Caribbeanislanders to make them acquainted with the richness andvalue of their dry shrublands. Preparation of a popularguide for regional use to the succulent plant species of the

West Indies would be useful. These programmes shouldaim to dispel the general misconception that the WestIndian thorn scrubs are useless, and to move people to theforefront of the battle to conserve the rare and uniqueplants that are only found in this type of vegetation;people should believe in developing different sustainableapproaches to the dry shrubland use, but only if this isaccomplished by evaluating the interaction between man-made and natural environments, and the ways in whichthis sets limits for social and economic aspects of life.

Contact: NYBG, MNHN, CIAC, CEA, NTCI

South America

92) B situ conservation. Organisation and help toregional botanic gardens as well as germplasm centrcs isneeded for future action.

93) Taxonomic research. Provide data and facilitiesto those who are currently revising the taxonomy ofsucculent plants. Encourage those to include informationon the conservation status of the species treated.

94) Study of the South American cactus trade.Investigation of the collection and propagation of SouthAmerican cacti for international trade, including trade inwild-collected seed, focusing on nurseries and commercialdealers within the region. Investigation of trade patternswithin South America concentrating on Peru, Chile andBrazil.

Contact: TRAFFIC South America

95) Field surveys of the taxonomic andconservation status of Parodia (Notocactus), Frailea,and Gymnocalycium. The conservation status of most ofthe Parodia (Notocactus) taxa (many of doubtfultaxonomic standing), c. 15 Frailea and 6 Gyr7z?zocalyC’ir1171spp. (2 endemic), in the southern part of Rio Grande doSul, Brazil and Uruguay needs to be determined. Somepopulations are known to be very small and illegalcollection to satisfy the demand for novelties by hobbyistsin Europe and elsewhere is certainly taking place. Fieldsurveys are needed to assess the taxonomic andconservation status of these taxa and to investigate thepossibilities for in situ conservation.

Colombia/Venezuela96) Assessment of conservation status. Fieldsurveys are needed to assess the conservation status oflocal species with restricted distributions in northernVenezuela and Colombia: Agave cob, Echevel-ia sp. nov.(Lara state), Cereus fkicii (C. nissellianus), C. mortemwii,C. horrispinus a l l f r o m n o r t h e r n Venezuela;Pseudoacanthocereus (Acan thocereus) siccri-isrlcrlsis.northern Venezuela and Colombia; Amatocer-eus hrtr?lilis,Rio Dagua valley, western Colombia.

142

Venezuela97) Creation of a reserve. Consideration should begiven to setting aside a site within the Venezuelan rangeof Melocactus schatzlii, e.g. south of Ejido, which couldalso protect a currently undescribed endemic Echeveriaspecies.

Ecuador

98) Survey of Catamayo valley. Determine status ofall succulent species in Catamayo valley: Armatocereusbrevispinus, Cleistocactus leonensis, and Espostoafkutescens are rare and restricted to the valley as are someother, non-endemic succulents and Weberocereus rosei,known from only two natural sites in Chimborazo andCaoar Provinces.

Peru99) Assessment of the in situ c o n s e r v a t i o nrequirements of succulents. Assessment of the extentto which the protected area system of Peru protects thehabitats of endemic succulents. The Andean Region ofPeru is extremely rich in succulent plant species, with highlevels of endemism. Survey work is needed throughoutthe drier areas. In the Coastal Region, surveys areneeded to determine the degree of threat to populationsof particular succulent species in the more mesic areas.

100) Peru/Bolivian Andes. Field study is needed toresolve questions of taxonomic and conservation statusconsidering the high endemicity in this region.

Bolivia101) j!Tx situ conservation. Support is needed toencourage the ex situ conservation of threatened cacti inthe newly formed botanic garden at the University of LaPaz.

Chile102) I n situ protection. The followingrecommendations for protected areas result from fieldwork carried out by the SSC Cactus and Succulent Group(Anderson et al. 1990):

a) Emphasise the establishment of Paposo NationalPark.

b) Expand the boundaries of Pan de Azucar NationalPark. It is important to include the completeranges of the very restricted taxa Copiapoa Maui, C.longistaminea, and C. sevpentisulcata within thePark’s protection. If possible the Park shouldextend north to Taltal and south to Chanaral, thenorthern extension including the entire ranges ofthe rare Copiapoa rupestris and C. desertorum.

c) Establish a national park or protected areabetween Totoral and Huasco. Such a legally

d)

e>

protected zone would specifically prescrvc thespectacular and remarkable populations ofCopiapoa malletiana.Set aside Quedrada El Leon as a nature reserve.This canyon, located near the town of Caldcra,contains an important mixture of the northern andsouthern cactus and other floristic elements.Expand the boundary of the proposed wetlandsnature reserve to include Punta de Teatinos, whichis north of La Serena.

103) Survey of Coastal Fog Desert. Further study ofthe endemic flora and ecology of the Coastal Fog Desertand field surveys to determine range and conservationstatus of taxa affected by mining and ore processing in thevicinity of Tocopilla and Copiapo.

104) Evaluation of the conservation status of theChilean Cactaceae. A comprehensive proposal to studythe taxonomy, distribution, ecology, and conservationstatus of all Chilean cacti was presented at the Sixthmeeting of the CITES Plants Committee. This wouldform the basis for implementation and improvement ofconservation legislation and regulation of the uses of cactiresources.

Contact : Departamento de Protection de 10sRecursos Naturales Renovables (Ricardo Scheu)Budget: US$334,903

Brazil105) Creation of reserves in eastern Brazil. Themost serious conservation problem in Brazil at present isthe almost complete lack of federally-run rcscrvcs toprotect the rare and endemic terrestrial succulents fromthe dry parts of north-east and south-east Brazil, and fromthe rocky east Brazilian Highlands, which rise out of thedry zone.

a>

b)

C>

Rio Jequitinhonha valley, middle section (i.c.Aracuai to Jacinto), north-eastern Minas Gcrais.One of the most important areas needingprotection amongst the southern caatinga-agrcstcvegetation zone where a remarkably richassortment of endemic and potentially threatenedcactus species exists.Rio de Contas, Mun. Maracas, Bahia, just cast ofPorto Alegre on the north bank. Here lies acomprehensive range of southern caatinga cacti,including the rare Espostoopsis dybowkyi.S5o Francisco River valley with its deep soils and

Bambui limestone outcrops in the (especially for

Bombacaceae and columnar Cactaceae) hosts twopotential sites:i) massive raised outcrop south of the town of

Iuiu on the east bank of the river (Bahia)hosting some spectacular bottle-trees ofCavaniZZesia and Ceiba as well as two very local

143

ii)

endemics (Facheiroa estevesii, Opuntia estevesii)restricted to the rock itself,west side of the river where further endemics,such as the Micranthocereus dolichosperma ticusand Facheiroa cephaliomelana, are located.

Locations where new protected areas are needed toassist the conservation of the rare cacti of eastern Brazil,including the CITES Appendix I cactus taxa, are asfollows:

49

4

4

g>h)9.J>

k)

1)

m)

Serra Dourada (Go&),arenitic outcrops 20-25 km west of Morro doChap& (Bahia),southern end of the Serra Chapada and associatedcerrado (27-28 km W of Seabra, BA),quartzitic outcrops at Brejinho das Ametistas (S

BA)Se&a Geral c. 12-15 km east of Monte Azul(Minas Gerais),Serra Geral with white sand cerrado 12 km east ofMato Verde (MG),Serra Bocaina (N of Grao Mogol, MG),cerrado and mountains around Grao Mogol itself,Serra do Cabral (MG),western slopes of the Serra de Minas east of SantaBarbara (Mun. August0 de Lima, MG),one or more sites in the vicinity of Diamantina

(MG),Serra Negra between Itamarandiba and RioVermelho (MG),Serra do Caraca (Mun. Santa Barbara, MG)

Extens ion o f the Parque National ChapadaDiamantina, Bahia slightly to its west, to include a secondpopulation of the remarkable Arrojada bahiensis.

Field survey to investigate the feasibility ofrecommending protected areas for rare and probablyvulnerable (but inadequately studied) species ofRhipsalideae from the region of Rio de Janeiro such asRhipsalis cereoides, R. mesembryanthemoides, andSchlumbergera orssichiana.

106) Monitoring of epiphytic cacti in the Atlantic CoastalForest. Monitoring by local scientists of the status andlevel of protection afforded to epiphytic cacti at thefollowing sites:

a) Parques Nacionais da Floresta da Tijuca and daSerra dos Orgaos (RJ),

b) Parques Estaduais de Ilha Grande (RJ), deIbitipoca (MG), de Campos do Jordao and dePicinguaba (SP),

c) Reservas Biologicas de Polo das Antas, deParanapiacaba, da Jureia, Ilhabela, and thatproposed for the Serra do Japi (SP).

Paraguay107) Designation of Cerro Le6n as a national park.

108) Field surveys in the Cordillera de 10s Altos todetermine status of smaller-growing species.

144

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Annex 1

Members of the Agavaceae withrestricted distribution’

Prepared by Wendy Hodgson and Abisai Garcia Mendoza.

Conservation status

CITES FWP

IUCN3 SEMARNAPS

Agave acicularis Trek.A. acklinicola Trek.A. aktites GentryA. angustifolia Haw. var.

nivea (Trel.) GentryA. arizonica Gentry

& J.WeberA. asperrima Jacobi ssp.

maderensis (Gentry) UllrichA. asperrima ssp.

potosiensis (Gentry) UllrichA. asperrima ssp.

zarcensis (Gentry) UllrichA. atrovirens Karw. var.

mirabilis (Trel.) GentryA. attenuata Salm-DyckA. avellanidens Trel.A. bahamana Trel.A. braceana Trel.A. bracteosa S.Watson

ex Engelm.A. cacozela Trel.A. capensis GentryA. celsii Hook.f. var.

albicans (Jacobi) GentryA. chiapensis JacobiA. chrysoglossa I.M.JohnstonA. colimana GentryA. congesta GentryA. cupreata Trel. & BergerA. dasylirioides Jacobi &

BoucheA. deamiana Trel.A. eggersiana Trel.A. felgeri GentryA. filifera Salm-DyckA. fortiflora GentryA. funkiana Koch & BoucheA. geminiflora (Tagl.)

Ker GawlerA. gigantensis GentryA. glomerulifiora (Engelm.)

BergerA. grisea Trel.A. guiengola GentryA. gypsophila GentryA. hiemiflora GentryA. hookeri JacobiA. horrida Lem. ex JacobiA. hurteri Trel.A. impressa GentryA. inaguensis Trel.

EI

R

I E E

A

R

R

A

I c2EV A

R

ntV A

Conservation statusCITES FWSJ

IUCNJ SEMARNAP-5

A. inaequidens Koch ssp.barrancensis Gentry

A. indagatoncm Trel.A. inter-m&a Trel.A. isthmensis Garcia-Mend. & PalmaA. jaibolie GentryA. kellermaniana Trel.A. kewensis Jacobi RA. lagunae Trel. RA. longipes Trel. RA. lurida Aiton EA. margaritae BrandegeeA. mckelveyana Gentry C3A. millspaughii Trel. I

A. minarum Trel.A. moranii GentryA. murpheyi Gibson R c2A. nashii Trel.A. nayaritensis GentryA. neglecta SmallA. nizandensis Cutak EA. obscura SchiedeA. ocahui Gentry

var. ocahuiA. ocahui

var. longifolia GentryA. ornithobroma GentryA. oroensis GentryA. panamana Trel.A. papyrocarpa Trel. EA. parrasana Berger R RA. parviflora

ssp. flexiflora Gentry I v AA. parviflora Torr.

ssp. parviflora I R C2 RA. peacockii Croucher R RA. pelona GentryA. pendula Schnittsp.A. polian thiflora AA. potrerana Trel.A. promontori Trel.A. pumila De Smet ex BakerA. rhodacantha Trel.A. scaposa GentryA. schottii Engelm. var.

treleasei (Tourney)Kearney & Peebles E C2

A. sebastiana GreeneA. shrevei Gentry

ssp. matapensis GentryA. sobria Brandegee ssp.

frailensis Gentry

156

Conservation status

CITES FWS4

IUCNJ SEMARNAPS

A. sobria ssp. roseana (Trel.) GentryA. stringens Trel.A. subsimplex Trel.A. tecta Trel.A. thomasae Trel.A. titanota GentryA. triangularis JacobiA. utahensis Engelm. var.

eborispina (Hes.) BreitungA. victoriae-reginae MooreA. vizcainoensis GentryA. warelliana BakerA. weberi Cels ex Poiss.A. wendtii Basanez, sp. nov. in ed.A. wercklei Weber ex BergerA. zebra GentryAgave sp. (central Arizona)Beaucarnea congesta Hernandez,

sp. nov. in ed.B. goldmanii RoseB. gracihs LemaireB. guaternalensis RoseB. hiriartiae HernandezB. pliabilis (Baker) RoseB. purpusii LemaireB. recurvata Lemaire (incl. B.

inermis [S.Watson] Rose)B. stricta LemaireBeschorneria albiflora

MatudaB. calcicola Garcia-Mend.B. tubiflora (Kunth & Bouche)

KunthB. wrightii Hook.Calibanus hookeri (Lem.) Trel.Dasylirion longissimumD. longistylum Ma&rideD. palaciosii Rzed.Furcraea bedinghausii KochF. cabuya Trel.F. cahum Trel.F. guatemalensis Trel.F. guerrerensis MatudaF. hexapetala (Jacq.) UrbanF. macdougalii MatudaF. macrophylla BakerF. melanodonta Trel.F. stratiotes Boye-PedersenF. tuberosa (Willd.) W.T.AitonHesperaloe chiangii Starr,

sp. nov. in ed.H. finifera (Koch) Trel.H. nocturna GentryH. parviflora (Torr.) CoulterManfredae brunneaManjreda chamelensis Lott

& VerhoekM. elongata RoseA4. fusca RavennaA4. guerrerensis MatudaM. hauniensis (Boyc-Petersen)

Verhoek

R RR RR R

nt c3II E P

R R

E

c 2

AA

AAAA

A

RR

RI

V

I

R

R

R

RR

R

A

RA

P

Conservation status

CITES FWV

IUCNJ SEMARNAP’

M. involuta McVaughM. longibracteata VerhoekM. longiflora (Rose) Verhoek AM. maculata (Mart.) RoseM. nanchititlensis Matuda AM. planifolia (S.Watson) RoseM. potosina (Robinson &

Greenm.) RoseM. pubescens (Regel & Ortega)

Verhoek ex I.L.PinaM. revoluta (Klotsch) RoseM. rubescens RoseM. sileri VerhoekNolina arenicola CorrellN. atopocarpa BartlettN. beldingii Brandegee var.

deserticola Trel.N. brittoniana Nash EN. cespitifera Trel.N. elegans RoseN. humilis S.WatsonN. inter-rata Gentry I EN. lindheimeriana

(KSchum.) S.WatsonN. nelsoni RoseN. palmeri S.Watson var. palmeriN. palmeri var. brandegeei Trel.N. pumila RoseN. parryi S.Watson var. wolfii MunzPleomele angustifolia N.E.Br.P. aurea (Mann.) N.E.Br. ntP. auwahiensis St.John ntP. elliptica (Thunb.) O.Deg.P. femaldii St.John ntP. flexuosa (Regel) O.Deg.P. forbesii O.Deg. ntP. halapepe St.John ntP. hawaiiensis O.Deg. & I.Deg.Pleomele sp. (Hawaii)Polianthes densiflora (Robinson R

& Fernald) ShinnerP. elongata RoseP. geminiflora (Lex.) Rose

var. clivicola McVaughP. gracilis Link & OttoP. howardii Verhoek RP. longiflora Rose RP. montana RoseP. nelsoni RoseP. palustris Rose RP. platyphylla Rose RP. sessiliflora (Hemsl.) RoseYucca angustissima Engelm.

& Trel. var. avia RevealY angustissima var. toftiae

(Welsh) Reveal EY. arizonica McKelvcy ntY. campestris McKelveyI’. coahuilensis Matuda & I.L.PinaY. elata Engelm. var. verdiensis

(McKelvey) RevealY. endlichiana Trel.

Conservation statusCITES FWP

IUCN” SEMARNAPS

Y grandiflora GentryY jaliscensis Trel.Y lacandonica Gbmez Pompa

& Valdez

Y. madrensis GentryY queretaroensisY necopina Shinner

Y potosina Rzed.Y queretaroensis 1.L Pina

Y. whipplei Torr. var.eremica Epling & Haines

R RVV A

ntR

E

1 Taxa considered to be micro-aerial (narrow) endemics inhabiting zones of less than 1OOkm in length and width.

2 CITES:

Appendix 1 No international trade allowed

Appendix II International trade only by permit

3 IUCN Categories at the global level (old categories of threat are used here): see Annex 16

4 US Fish and Wildlife Service (USFWS) listing:

E = Endangered: any species in danger of extinction throughout all or a significant portion of its range.

T = Threatened: any species likely to become an endangered species within the foreseeable future throughout all or a significant portion of its range.

Cl = Category 1 status: taxa for which the FWS has sufficient (but not necessarily complete) information on vulnerability and threats to support a proposal to

list them as threatened or endangered.

C2= Category 2 status: taxa for which the FWS has insufficient information to support a proposed rule to add the species to the threatened or endangered

species list; further biological research and field study will usually be needed to change the status of taxa in category 2.

C3= Category 3 status: taxa that are no longer being considered for listing as endangered or threatened (and included in one of the three subcategories).

5 SEMARNAP Categories for Mexico

A = threatened (amenazada)

R = rare (rara)

P = in danger of extinction (en peligro de extincion)

158

Annex 2

Asclepiadaceae of conservation concernPrepared bv Focke Albers and Ulrich Meve.

J

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Brachystelma S i m sB. alpinrm R.A.DyerB. arenarium SMooreB. attenuatum (Wight) Hook.B. australe R.A.DyerB. hlephuranthem HuberB. houmeae GambleB. brevipedicellutum TurrillB. hrevituhulutum (Bedd.) GambleB. buchananii N.E.Br.B. cuflrum (Schltr.) N.E.Br.B. campanulutum N.E.Br.B. canum R.A.DyerB. cathcartense R.A.DyerB. caudatum (Thunb.) N.E.Br.B. chlorozonum E.A.BruceB. ciliatum Arekal & RamakrishnaB. comptum N.E.Br.B. constrictum J .HallB. decipiens N.E.Br.B. delicatum R.A.DyerB. dimorphurn R.A.Dyer ssp. dimorphurnB. dimorphurn R.A.Dyer ssp. gratum R.A.DyerB. discoidcum R.A.DyerB. duplicatum R.A. DyerB. edulis Coll. & Hemsl.B. elegantulum S.MooreB. elenanduensis M.CharB. elongutum (Schltr.) N.E.Br.B. exile Bull.B. festuc(folium E.A.BruceB. furcatum BoeleB. gemmeum R.A.DycrB. gerrurdii Harv.B. glabr$orum (F.Muell.) Schltr.B. glabrum Hook.f.B. glenense R. A. DyerB. gracillimum R.A.DyerB. huttonii (Ha-v.) N.E.Br.B. incanum R.A.DyerB. keniense Schweinf.B. kerrii CraibB. kolarensisB. laevigatum (Wight) Hook.f.B. lancasteri BocleB. lunkana Dassan. & Jayag.B. letestui Peller.B. longifolium (Schltr.) N.E.Br.B. macropetalr~m (Schltr.) N.E.Br.B. maculatum Hook.f.B. medusanthemum J.-P.Lebrun & StorkB. merrillii Schltr.B. meyerianum Schltr.B. micranthum E.Mey.B. minimum R.A.DyerB. minor E.A.BruceB. modestum R.A.Dyer

159

ERE(?Ex)EI (?Ex)RIRRIKRRRERRRREEI (?Ex)RRREREExRRRRR

KKRRRRRRRVEKRKEERRRRRRRR

B. montanum R.A.DyerB. mortonii WalkerB. nutalense (Schltr.) N.E.Br.B. ngomense R.A.DyerB. occidentale Schltr.B. omissum Bull.B. pachypodium R.A.DyerB. papuanum Schltr.B. parviflorum (Wight) Hook.f.B. pan&m R.A.DyerB. pauciflorum DuthieB. perditum R.A.DyerB. petraeunz R.A.DyerB. pilosum R.A.DyerB. praelongum S.MooreB. prostratum E.A.BruceB. ramosissimum (Schltr.) N.E.Br.B. mngacharii GambleB. sandersonii (Oliv.) N.E.Br.B. schinzii (K.Schum.) N.E.Br.B. schizoglossoides (Schltr.) N.E.Br.B. sclzoenlalzdiaraun? Schltr.B. simplex Schltr.B. subaphyllum K.Schum.B. swazicum R.A.DyerB. tabulurium R.A.DyerB. tavulla K.Schum.B. tenellum R.A.DyerB. tenue R.A.DyerB. thunbergii N.E.Br.B. tuberosum R.Br.B. vahrmeijeri R.A.DyerB. villosum (Schltr.) N.E.Br.B. volubile Hook.f.

Caralluma R.Br.C. adscendens (Roxb.) R.Br. var. NIISUC~~~~CI~SC. arabica N.E.Br.C. baradii LavranosC. beviloba (P.R.O.Bally) M.G.GilbertC. bhupinderana SarkariaC. circes M.G.GilbertC. congestiflora P.R.O.BallyC. crenulata Wall.C. dicapuae (Chiov.) Chiov. ssp. dicllpllrrc

C. difidsa (Wight) N.E.Br.C. dodsoniana LavranosC. dolichocarpa SchwartzC. edwardsae (M.G.Gilbert) M.G.GilbertC. foetida BruceC. jkrta P.R.O.BallyC. joannis MaireC. lavruni Rauh & WertelC. laticorona (M.G.Gilbert) M.G.GilbertC. long$ora M.G.GilbertC. mireillac LavranosC. moniliformis P.R.O.BallyC. munbyana (Decne) N.E.Br. var. I?U&~YU~UC. nilugiriuna Kumani Rc RaoC. peckii P.R.O.Bally

RRRRRKKRRRKRKRR

RREExEREKRIERIERKRRKRKKVRKKEVRKEKRRRRERREERRRRRRKK

C. munbyana (Decne) N.E.Br. var. munbyanaC. nilagiriana Kumani & RaoC. peckii P.R.O.BallyC. peschii NelC. priogonium KSchum.C. procumbens Grav. & May.C. rauhii LavranosC. sarkariae Lavranos & FrandsenC. socrotrana (Ba1f.f.) N.E.Br.C. solenophora LavranosC. somalica N.E.Br.C. staintonii HaraC. truncatocoronata (Sedgw.) Grav. & May.C. umbellata Haw.C. vaduliae Lavranos

Ceropegia L.C. affinis VatkeC. albisepta Jum. & H.PerrierC. ampliata var. madagascariensis Lavranos & MoratC. antennifera Schltr.C. arenaria R.A.DyerC. aridicola W.W.Sm.C. armandii RauhC. arnottiana WightC. attenuataC. barbata R.A.DyerC. bamesii Bruce & ChatterjeeC. beddomei Hook.f.C. bhutanica HaraC. bosseri Rauh & BuchlohC. botrys K.Schum.C. bowkeri Harv. ssp. bowkeriC. bowkeri ssp. S.O. sororia (Harv. ex Hook.f.) R.A.DyerC. brevirostris P.R.O.Bally & FieldC. campanulata Don var. campanulataC. cancellata Rchb.C. candelabrum L. ssp. S.O. candelabrumC. ca taphyllaris Bull.C. cera tophora Sven t .C. chipiaensis Stopp.C. chortophylla Werderm.C. christenseniana Hand.-Mazz.C. chrysanthaC. cimiciodora Oberm.C. conrathii Schltr.C. convoluloides A.Rich.C. cycniflora R.A.DyerC. damannii Stopp.C. decidua Bruce ssp. deciduaC. decidua ssp. pretoriensisC. deightonii Hutch. & Dalzell ssp. deightoniiC. dimorpha HumbertC. dinteri Schltr.C. distincta ssp. vermcolosaC. dorjei C.E.C.Fisch.C. evansii McCannC. fantastica Sedgw.C. filiformis (Burch.) Schltr.C. fimbriata E.Mey. ssp. fimbriataC. fimbriata ssp. connivens (R.A.Dyer) BruynsC. fimbriata ssp. geniculata (R.A.Dyer) BruynsC. fimbrifera BeddomeC. floribunda N.E.Br.C. furcata Werderm.C. gemmifera K.Schum.

160

KKEERKKERRVRREKERKKRRKREKRERKKKKRKRRRVEEEEKRIKEKRREKKRVEKRRRRKKEKK

C. gilgiana Werderm.C. hirsuta Wight & Arn.C. hofstaetteri RauhC. huberi AnsariC. humbertii HuberC. illegitima HuberC. infZata Hochst. ex Chiov.C. insignis R.A.DyerC. jainii Ansari & B.G.P. Kulk.C. kachinensis PrainC. krainzii Svent.C. kundelunguensis MalaisseC. langkawiensis RintzC. lawii Hook.fC. ledermanii Schltr.C. leroyi Rauh & Marn.- Lap.C. lindenii LavranosC. loranthiflora K.Schum.C. ludlowii HuberC. maccannii AnsariC. madagascariensis Decne.C. madens Werderm.C. mafekingensis (N.E.Br.) R.A.DyerC. mahabalei Hem. & AnsariC. mairei (Lev.) Huber var. maireiC. maiuscula HuberC. mayottae HuberC. media (Huber) AnsariC. mendesii StoppC. mirabilis HuberC. monticola W.W.Sm.C. muliensis W.W.Sm.C. muzhingana MalaisseC. nana Coll. & Hemsl.C. noorjahanae AnsariC. ngoyana MalaisseC. nuda Hutch. & BruceC. occidentalis R.A.DyerC. occulta R.A.DyerC. odorata Nimmo ex Hook.f.C. omissa HuberC. panchganiensis Blatt. & McCannC. paohsingensos Tsiang & LiC. paricyma N.E.Br.C. parviflora TrimenC. peteri Werderm.C. petignatii RauhC. porphyrotricha W.W.Sm.C. pusilla Wight & Arn.C. radicans Schltr. ssp. radicansC. razafindratsirana (Rauh & Buchloh) RauhC. renzii StoppC. ringens A.RichC. rollae HemadriC. rudatisii Schltr.C. sahyadrica Ansari & Kulk.C. salicifolia HuberC. santapaui Wadhwa & AnsariC. saxatilis Jum. & Perr.C. scabra Jum. & Perr.C. scabyriflora N.E.Br.C. schliebenii Markgr.C. senegalensis HuberC. simoneae RauhC. sootepensis HuberC. speciosa Huber

RERRKKRRRRKKERK

KKKKKKKKKKKRKKKRKKKKKKKKKKRKKKKKRKKKKRKKEKKK

EER

C. spiralis WightC. stentiae BruceC. swaziorum D.V.FieldC. taprobanica HuberC. teniana Hand.-Mazz.C. thwaitesii HookC. tihamana Chaudhary & LavranosC. tomentosa Schltr.C. turricula BruceC. ugeni C.E.C.Fisch.C. vanderystii De Wild.C. vignaldiana A.Rich.C. vincaefolia HookC. viridis ChouxC. wallichii Wight

Cynanchum L.C. aequilongum ChouxC. ambositrense ChouxC. ampanihense Jum. & H.PerrierC. andringitrense ChouxC. angkavokeliense ChouxC. antandroy Desc.C. appendicula turn ChouxC. arenarium Jum. & H.PerrierC. bekinolense ChouxC. bisinuatum Jum. & H.PerrierC. bojerianum Decne.C. compactum ChouxC. cucullatum N.E.Br.C. danguyanum ChouxC. decaisneanum Desc.C. descii RauhC. fibriatum ChouxC. gerrerdii (Harvey) LiedeC. implicatum (Jum. & H.Perrier) Jum. & H.PerrierC. juliani-marnieri Dcsc.C. Jumi ChouxC. junc&$orme (Decne.)LiedeC. lecontei ChouxC. lineare N.E.Br.C. luteifluens (Jum. & H.Perrier) Desc.C. macranthum Jum. & H.PerrierC. macrolobum Jum. & H.PerrierC. madagascariense K.Schum.C. madecassum Desc.C. mahafalense Jum. & H.PerrierC. menarandrense Jum. & H.PerrierC. messeri (Buchenau) Jum. & H.PerrierC. marnieranum RauhC. moramangense ChouxC. napiferum ChouxC. napif~~rme ChouxC. nodosum (Jum. & H.Perrier) Desc.C. pachylobum ChouxC. papillatum ChouxC. radiatum Jum. & H.PerrierC. rossii RauhC. sessiliflorum (Decne.)LiedeC. subtilis LiedeC. surprisum Liede

Duvalia N.E.Br.

D. anemoniflora (Deflers) R.A.Dyer & LavranosD. galgallensis LavranosD. parviflora N.E.Br.

161

KR

E

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E

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D. pillansii N.E.Br.D. somalensis Lavranos

Duvaliandra M.G.GilbertD. dioscoriodes (Lavranos) M.G.Gilber

Echidnopsis Hook.f.E. angustiloba Bruce & P.R.O.BallyE. archeri P.R.O.BallyE. ballyi (Marnier-Lap.) P.R.O.BallyE. bihendulensis P.R.O.BallyE. ciliata P.R.O.BallyE. ericiflora LavranosE. insularis LavranosE. leachii LavranosE. malum (Lavranos) BruynsE. mijerteina Lavranos var. mijerteinaE. milleri LavranosE. montana (R.A.Dyer & Bruce) P.R.O.E. repens R.A.Dyer & I.Verd.E. seibanica LavranosE. socotrana LavranosE. squamulata (Decne.) P.R.O.BallyE. urceolata P.R.O.BallyE. virchowii K.Schum.

FolotsiaF. aculeatum Jum. & H.PerrierF. floribundum Desc.F. grandiflorum Jum. & H.PerrierF. madagascariense (Jum. & H.Perrier)F. sarcostemmoides cost. & Bois

Frerea DalzellF. indica Dalzell

Hoodia Sweet.H. dregei N.E.Br.H. juttae DinterH. mossamedensis (Leach) PlowesH. ojficinalis ssp. delaetiana (Dinter) BryunsH. pilifera (L.f.) Plowes ssp. annulata (N.E.Br.) BryunsH. pilifera ssp. pilifera (L.f.) PlowesH. pilifera ssp. pillansii (N.E.Br.) BryunsH. ruschii DinterH. triebneri (Nel) Bruyns

Huernia R.Br.H. andreaeana (Rauh) LeachH. arabica N.E.Br.H. archeri LeachH. baveri LeachH. boleana M.G.GilbertH. coninna N.E.Br.H. erectiloba Leach & LavranosH. erinacea P.R.O.BallyH. hadhramautica LavranosH. hallii E. & B.M. LambH. humilis (Masson) Haw.H. hystrti (Hook.f.)N.E.Br. var. panwllr L.C.LeachH. kennedyana LavranosH. lea&ii LavranosH. lodarensis LavranosH. longii PillansH. marnieriana LavranosH. nigeriana Lavranos

VIRRRRRRIIRExRREx

R

E

R

R

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H. nouhuysii I.Verd.H. pendulu E.A.BruceH. piersii N.E.Br.H. plowesii LeachH. praestans N.E.Br.H. procumbens (R.A.Dyer) LeachH. quinta (Phillips) A.C. White & B. SloaneH. recondita M.G.GilbertH. schneideriana BergerH. similis N.E.Br.H. tanganyikensis Bruce & P.R.O.BallyH. thudichumii LeachH. urceolata LeachH. whitesloaneana NelH. witzenbergensis C.A. Liickh.

Huerniopsis N.E.Br.H. atrosanguinea (N.E.Br.) A.C. Wight & B.Sloane

Karimbolea

Karimbloea verrucosa Desc.

Lavrania Plowes

L. haagnerae Plowes

Notechidnopsis Lavranos & BleckN. columnaris (Nel) Lavranos & Bleck

Orbea (L.) Haw.

0. ciliata (Thunb.) Leach0. halipedicola Leach ssp. halipedicola0. macloughlinii (I.Verd.) Leach0. paradoxa (I.Verd.) Leach0. prognatha (P.R.O.Bally) Leach0. rangeana (Dinter & Berger) Leach0. speciosa Leach0. woodii (N.E.Br.) Leach

Orbeanthus Leach

0. conjunctus (A.C. White & B. Sloane) Leach0. hardyi (R.A.Dyer) Leach

Orbeopsis Leach0. albocastanea (Marloth) Leach0. gerstneri (Letty) Leach ssp. elongata (R.A.Dyer) Leach0. gerstneri ssp. gerstneri0. gossweileri (S.Moore) Leach0. huillensis (Hiern) Leach0. knobelii (Phillips) Leach0. tsumebensis (Oberm.) Leach

Pachycymbium Leach

P. abayense (M.G.Gilbert) M.G.GilbertP. araysianum (Lavranos & Bilaidi) M.G.GilbertP. chlysostephanum (Deflers) M.G.GilbertP. denboefii (Lavranos) M.G.GilbertP. distinctum (E.A.Bruce) M.G.GilbertP. eremastrum (Schwartz) M.G.GilbertP. gemugofanum (M.G.Gilbert) M.G.GilbertP. huernioides (P.R.O.Bally) M.G.GilbertP. kochii (Lavranos) M.G.GilbertP. laikipiense M.G.GilbertP. lancasteri LavranosP. meintjesianum (Lavranos) M.G.GilbertP. rogersii (L.Bolus) M.G.GilbertP. sacculatum (N.E.Br.) M.G.GilbertP. sprengeri ssp. ogadense (M.G.Gilbert) M-G-Gilbert

162

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P. tubiforme (Bruce & P.R.O.Bally) M.G.GilbertP. umbomboense (I.Verd.) M.G.GilbertP. wilsonii (P.R.O.Bally) M.G.GilbertPectinaria HawP. articulata (Aiton) Haw. ssp. articulataP. articulata ssp. borealis BruynsP. longipes (N.E.Br.) Bruyns

Piaranthus R.Br.P. barrydalensis MeveP. fiamesii Pillans

Pseudolithos P.R.0 Bally

P. caput-viperae LavranosP. cubiformis (P.R.O.Bally) P.R.O.BallyP. horwoodii P.R.O.Bally & LavranosP. migiurtinus (Chiov.) P.R.O.Bally

Quaqua N.E.Br.Q. arrnata (N.E.Br.) Bruyns ssp. arenicola (N.E.Br.) BruynsQ. armata ssp. maritima BruynsQ. framesii (Pillans) BruynsQ. inversa (N.E.Br.) Bruyns var. cincta (C.A. Luckh.) BruynsQ. inversa (N.E.Br.) Bruyns ssp. inversaQ. linearis (N.E.Br.) BruynsQ. multiflora (R.A.Dyer) BruynsQ. parviflora ssp. bayeriana BruynsQ. pruinosa (Masson) Bruyns

Rhytidocaulon P.R.O.BallyR. fulleri Lavranos & Mort.R. paradoxum P.R.O.BallyR. sheilae FieldR. subscandens P.R.O.BallyR. tortum (N.R.Br.) M.G.GilbertR. richardianum Lavranos

Riocreuxia DecneR. aberrans R.A.DyerR. alexandrina R.A.DyerR. bolusii N.E.Br.R. chrysochroma (Huber) A.C. SmithR. nepalensis A.C.SmithR. splendida K.Schum.R. woodii N.E.Br.

Sarcostemma R.Br.S. decorsei Cost. & Gall.S. insignis N.E.Br.S. madagascariensis Desc.S. viminale R.Br.

Stapelia L.S. baylissii LeachS. cedrimontana FrandsenS. clavicorona I.Verd.S. divaricata MassonS. erectiflora N.E.Br. var. prostrutiflora L.C. LeachS. glabricaulis N.E.Br.S. immelmaniae PillansS. kouga bergensis LeachS. montana Leach var. montanaS. obducta LeachS. paniculata Willd.S. parvula Kers.S. pearsonii N.E.Br.S. peglerae N.E.Br.

RRR/VR/VWVRE

RKRRRRRRR

VRERVR

S. pillansii N.E.Br. var. pillansiiS. praetermissa Leach var. praetermissa LeachS. remota R.A.DyerS. rubiginosa NelS. scitula LeachS. tsomoensis N.E.Br.S. unicornis C.A. Luckh.

Stapelianthus ChouxS. arenarius Bosser & MoratS. calcarophilus MoratS. decaryi ChouxS. hardyi LavranosS. insignis Desc. var. insignisS. insignis var. tangoboryensis RauhS. keraudreniae Bosser & MoratS. montagnacii (Boiteau) Boiteau &z BertrandS. pilosus (Choux) Lavranos & Hardy

Stapeliopsis Pillans

S. breviloba (R.A.Dyer) BruynsS. exasperata (Bruyns) BruynsS. neronis PillansS. pillansii (N.E.Br.) BruynsS. saxatilis N.E.Br. ssp. stayneri (M.B.Bayer) BruynsS. urniflora Lavranos

R

R

RRRKR/VRRWVRRR

Tavaresia Welw.T. angolensis WeIw.Tenaris E. MeyT. schultzei (Schltr.) Phillips

Tridentea Haw.

T. baylisii (Leach) var. baylisiiT. baylisii (Leach) var. ciliataT. choanantha (Lavranos & Hall) LeachT dwequensis (C.A.Luckh.) LeachT. herrei (Nel) LeachT. longii (C.A.Luckh.) LeachT. marientalensis ssp. albipilosa (Giess) LeachT. pachyrrhiza (Dinter) LeachT. parvipuncta var. truncata (C.A. Liickh.) LeachT. peculiaris (C.A.Luckh.) LeachT. ruschiana (Dinter) Leach

Tromotriche Haw.

T. engleriana (Schltr.) LeachT. revoluta (Masson) Haw.T. thudichumii (Pillans) Leac h

White-slonea Chiov.

W. crassa (N.E.Br.) Chiov.

’ Old TUCN Red List categories are used. See Annex 16 for definitions.

163

Annex 3

Succulents regulated by CITESSource: CITES/WCMC 1996.

AGAVACEAE

Agave arizonicaA. parvifloraA. victoriae-reginaeNolina in terra ta

APOCYNACEAE

Pachypodium entire genusP. am bongenseP. baroniiP. decaryiRauvolfia serpentina

ASCLEPIADACEAE

Ceropegia en tire genusFrerea indica

CACTACEAEEntire familvAriocarpus entire genusAstrophytum asteriasAztekium ritteriCoryphantha werderrnanniiDiscocactus entire genusDisocactus macdougallii

(Nopalxochia macdougallii)Echinocereus ferreirianus

var. lindsayiE. schmolliiEscobaria minima

(E. nellieae, Coryphantha minima)E. sneedii var. leei

- var. sneediiMammillaria pectiniferaM. solisioidesMelocactus conoideusM. deinacanthusM.M

glaucescenspaucispinus

Obregonia denegriiPachycereus militarisPediocactus bradyiP. despainiiP. kno wltoniiP. paradineiP. peeblesianusP. sileriP. winkleriPelecyphora entire genusSclerocactus brevihama tusS. erec tocen trus

(EchinomastusS. glaucus

erectocen trus)

S. mariposensisS. mesae-verdae

E.

S. papyracanthusS. pubispinusS. wrigh tiae

mariposensis)

CITESlisting’

IIIII

IIIIIII

IIII

IIIIIIII

I

II

IIIIIIIIIIIIIIIIIIII

IIIIII

Strombocactus disciformisTurbinicarpus entire genusUebelmannia entire genus

DIDIEREACEAEEntire family

EUPHORBIACEAE

Euphorbia entire genusE. ambovombensisE. cremersiiE. cylindrifoliaE. decaryiE. francoisiiE. moratiiE. parvicya thophoraE. quartziticolaE. tulearensis

FOUQUIERIACEAEFouquieria columnarisF. fascicula taF. purpusii

LILIACEAE

Aloe entire genus (except A. Vera)A. albidaA. albifloraA. alj?ediiA. bakeriA. bella tulaA. calcairophilaA. compressaA. delphinensisA. descoingsiiA. fiagilisA. ha worthioidesA. helenaeA. laetaA. parallelifoliaA. parvulaA. pillansiiA. polyphyllaA. rauhiiA. suzannaeA. thorncroftiiA. versicolorA. vossii

PORTULACACEAEAnacampseros entire genusCistanthe tweedyi (Lewisia tweedyi)Lewisia cotyledonL. maguireiL. serrata

‘Appendix I lists endangered species; international trade of wildspecimens is banned under the terms of the CITES Convention.Appendix II lists species which may be threatened by excessivelevels of trade and for which trade is permitted but controlledand monitored through a licensing system.

164

Annex 4

Succulents of Kenya of highest conservationconcern

Compiled by Len Newton, 1995.

AIZOACEAEDelosperma abyssinicum (Regel) Schwantes

ALOACEAEAloe archeri LavranosA. ballyi ReynoldsA. juvenna Brandham & S.CarterA. massawana ReynoldsA. microdonta ChiovendaA. parvidens M.G.Gilbert & SebsebeA. tugenensis L.E.Netwon & LavranosA. wrefordii Reynolds

ASCLEPIADACEAEBrachystelma lineare A.RichardCaralluma distincta BruceC. tubiformis Bruce & BallyC. vibratilis Bruce & BallyCeropegia albisepta Jum. & Perr. var. robynsiana H.HuberC. ampliata E.Meyer var. oqloba H.HuberC. ballyana BullockC. crassifolia Schlechter var. copleyae (Bruce & Bally) H.HuberC. galeata H.HuberC. somalensis ChiovendaC. stenoloba Hochst. ex Chiovenda var. moyalensis H.HuberC. variegata DecaisneEchidnopsis angustiloba Bruce & BallyE. ericiflora LavranosE. mariae LavranosE. radians BleckE. urceolata BallyHuernia andreaeana (Rauh) LeachH. archeri LeachH. keniensis R.E.Fries var. molonyae White & SloaneOrbea semota (N.E.Br.) LeachRhytidocaulon paradoxum Bally

CRASSULACEAEKalanchoe abyssinica ssp. hildebrandtiiK. aubrevillei CufodontisK. bipartita ChiovendaK. boranae RaadtsK. fadeniorum RaadtsK. obtusa Engler

CURCURBITACEAECephalopentandra ecirrhosa (Cogn.) Jeffrey

EUPHORBIACEAEEuphorbia ballyana RauhE. borenensis GilbertE. brevitorta BallyE. brunellii ChiovendaE. classenii Bally & S.CarterE. cussonioides BallyE. pseudoburuana Bally & S.CarterE. robecchii PaxE. tanaensis BallyE. taruensis SCarterE. turkanensis S.CarterE. wakefieldii N.E.Br.

Jatropha hildebrandtii Pax var. hildebrandtiiMonadenium rhizophonlm BallyM. rubellum (Bally) S.CarterM. stapelioides Pax var. congestum (Bally) S.CarterM. trinerve BallyM. yattanum Bally

ICACINACEAEPyrenacantha malvifolia Engler

MORACEAEDorstenia bamimiana Schweinfurth var. tropaeolifoliuD. zanzibarica Oliver

PASSIFLORACEAEAdenia globosa Engler ssp. globosa

PERIPLOCACEAERaphionacme madiensis S.Moore

PORTULACACEAEPortulaca ciferriiP. grandisP. oblongaP. pilosa

VITACEAECissus quadrangularis L. var. aculeatangula VerdcourtC. rotundifolia (Forsskal) Vahl var. .ferrugineu

165

Annex 5

Provisional list of succulent speciesof the Mediterranean Region

Based on text compiled by Silvio Fici and MauricioSajeva. Table compiled by M. Sajeva and Henk t’Hart.

The Mediterranean phytogeographic region is generallyconsidered to include the coastal fringe of theMediterranean area with the exception of parts of theLibyan, Egyptian, and Tunisian coasts, but with theinclusion of the Atlantic coasts of Portugal, Spain, andMorocco. Macaronesia (Canaries, Madeira, and theAzores) and parts of the Black Sea coasts have aMediterranean climate and are often included in thisphytogeographic region.

Within the region there is considerable geologicaland climatic diversity which is reflected in the vegetationand flora. In large areas of the region, however, thousandsof years of human influence, involving deforestation,overgrazing, fire, and urbanisation, has caused completemodification of the landscape and vegetation dynamics.

Succulent plants are relatively scarce in theMediterranean flora; those that do occur are usually smallin stature. They occur in various different habitat types,predominantly in montane regions. In contrast to thewidely dispersed Macaronisian succulent flora, theseplants are relatively dominant in only a few areas in theMediterranean basin, most notably on the Atlantic coastof Morocco. Some succulents, such as Caralluma andKalanchoe, belong to tropical or subtropical genera thatreach their northern limits in this region. Most succulents,however, belong to genera of the Crassulaceae which arewidespread in boreal regions, principally Sedum,Jovibarba, Sempervivum, and Rosularia. The succulentflora of Morocco has some genera in common with theCanary Islands, notably Aeonium, Aichryson, Caralluma,Euphorbia, and Kleinia, and a few species are common toboth.

The Atlantic coastal area of Morocco near Agadirand to the south where there are many succulents is apriority area for protection. Species include Euphorbiaoflicinarum, E. echinus, Caralluma burchardii, C. europaea,and Kleinia anteuphorbium. Threats to these plantsinclude industrial and tourism development, agriculture,and overgrazing.

The main threat for the survival of cacti andsucculents in this region is habitat modification anddestruction, mainly from industrial and tourismdevelopment of coastal and mountain grassland areas.

Caralluma europaea on Lampedusa Island.

Refuse disposal from construction also poses a threat tosome habitats, for example that of Caralluma europaea onLampedusa Island, Italy. Other threats to mountainoushabitats are dams and mining activities. Grazing,especially by goats, is a major factor in the loss ofbiodiversity in the Mediterranean region. Al though notserious, collection by individuals poses a threat to somesmall populations of rare mountain succulents within theregion. Fire destroys large areas of drier vegetation wherethese plants are often found. Invasive species, such asOpuntia &us-indica are a problem for native species inparts of Spain and Greece primarily threatening Aeonirmzarboreum and Dracaena draco, both native in north-westAfrica.

Existing conservation measuresEurope is generally well endowed with protected areas,but the Mediterranean region is probably that with theleast coverage. For the Mediterranean region the BernConvention only lists nine succulent species of the CanaryIslands. There are a few succulents from the region listedon CITES, but a significant trade in cacti and otherCITES succulents continues in the Mediterranean.Botanical gardens in the region which have importantsucculent collections include Blanes (Spain); TheAlmeda, Gibraltar Botanic Gardens; Monte Carlo(Monaco); and Palermo (Sicily).

166

Status1 Taxon Distribution3

E

V

IKntR

ntR

RV/R

R

AIZOACEAEAizoon canarienseA. hispanicumMesembryanthemum crystallinumM. gausseniiA4. nodifolium

ASCLEPIADACEAECaralluma aaronisC. burchardiiC. commutata ssp. hesperidiumC. europaea ssp. europaeaC. europaea ssp. maroccanaC. joannisC. munbyanaC. negevensisC. sinaicaC. tombuctuensisC. venenosa

COMPOSITAEKleinia anteuphorbium

CRASSULACEAEAeonium korneliuslemsiiHylotelephium anacampserosH. telephiumKalanchoe laciniataPistorinia brevifloraP. hispanicaRhodiola roseaRosularia aizoonR. blepharophyllaR. chrysanthaR. davisiiR. elyq’maiticuR. globularit$oliaR. haussknectiiR. hirsutaR. jaccardianaR. kesrouanensisR. lineataR. pallidifloraR. parvi$oliaR. radic$ora ssp. glabraR. radiciflora ssp. kurducaR. radiciflora SSQ. radicifloraR. sempewivum SSQ. sempervivumR. sempervivum SSQ. amanensisR. sempervivum SSQ. glaucophyllaR. sempervivum SSQ. kurdicaR. sempervivum SSQ. libanoticaR. sempervivum SSQ. persica:R. sempervivum SSQ. pestalozzaeR. serpentinica SSQ. serpentinicaR. serpentinica SSQ. giganteaR. serrataSedum aetnense

S. acreS. albumS. alpestreS. alsinefoliumS. amplexicaule ssp. amplexicaule

(southern Mediterranean)(widely distributed)(non perennial, widely distributed)

& (El(widely distributed)

Eg, IJMaMaSi, Eg, Li, Tn, AgHsMa (I)Hs (R), Ag, MaIJ, Sn

IJ, W Eg (E)AgAg (V>

Ma

MaHs, Ga, ItLu, Hs, Ga, Co, Sa, It, Ju, Al, Bu, RK, Gr, Tu, AnMaAg, Ma, TnHs, MaBu (R), Hs, Ga, It

An (1)An (only known from 2 localities)AnAn (R) (only known from 2 collections)AnAE, An (nt), Cy, LS

An CR)Ga, Hs, It, Lu, North AfricaMaLSLS, Jl, Sn

An, CYLSAn, LSAnAnAnAn, LSAnAn, LSAn, IJ, LS, Sn (R)An, LSAn, LSAnAnGr, Cr, AE, AnHs (V), Si (E), Ju, Al, Bu (R), RK, An; this species is a minute, ephemeral, undcrcollccted

annual; extremely local and rare throughout Europe, but quite common in AnatoliaLu, Hs, Ga, Si, It, Ju, Al, Bu, RK, Cr, AE, An, Li, Tn, Ag, MaLu, Hs, Bl, Ga, Co, Sa, Sl, It, Ju, Al, Bu, RK, Gr, Cr, Tu, An, LS, Li, Tn, Ag, MaIt, Ju, Gr, AnCo, It; quite local and rare in Liguria and PiemonteLu, Hs, Ga, ?Ma

167

VR

VRRR

R

Knt

R

R

RRRnt

RV

K

RR

ExntR

S. amplexicaule ssp. tenut$oliumS. andegavenseS. an&urnS. annuumS. apoleiponS. atratumS. arenariumS. assyriacumS. bractea turnS. brevifoliumS. caeruleumS. caricumS. cepaeaS. coespitosumS. confertiflorumS. creticum var. creticumS. creticum var. monocarpicumS. cypriumS. cyrenaicumS. dasyphyllumS. euxinumS. forsterianumS. fragransS. ga ttefosseiS. gracileS. grisebachii

S. gypsicolumS. hispanicumS. inconspicuumS. jahandieziiS. laconicum ssp. laconicumS. laconicum ssp. pentapolitanumS. laconicum ssp. pallidumS. lampusaeS. litoreumS. lydiumS. magellenseS. maw-urnS. melanantherumS. microstachyumS. modestumS. monregalenseS. montanumS. mucizoniaS. multicepsS. nanumS. ochroleucumS. palestinumS. pallidumS. pedicellatum ssp. lusitanicumS. pedicella turn ssp. pedicella turnS. pilosumS. polystriatumS. porphyreumS. pruina turn

S. pubescensS. obtusifoliumS. rubens

S. rupestreS. samium ssp. samiumS. schizolepsis

S. sediforme

AE, Al, An, Bu, Cr, Gr, It, Ju, Si, TuLu, Hs, Ga, Co, Sa, ItLu, Hs, GaAl, An, Bu, Co, Ga, Gr, Hs, It, Ju, TuGrHs, Ga, It, Ju, AL, Bu, Gr, AnLu, Hs; quite common and widely spread in large part of Central Spain and PortugalAn, LS; extremely local and rare, only in temporarily wet placesLi (R)Lu, Hs, Ga, Co, Sa, MaCo, Sa, Me, Si, Li, Tn, Ag, MaAE, An; confined to Rhodos and adjacent Anatolia; probably a ssp. of S. rubensHs, Ga, Co, Sa, Si, It, Ju, Al, Bu, Gr, AE, An, LS, Li, Tn, Agall but AL, Sn, EgAE (R), AnCr; restricted to east CreteCr

CY 0Li (R)all but RK, AE, Tu, Cy, LS, IJ, EG, Li; rare in AnatoliaAnGa, Hs, Ma, LuGa, It

Ma (R)AnJu, Al, Bu, Gr; comprises two ecotypes, of which the lowland form (up to c. I SO0 m) has two

cytotypes (2n= 16,32)Hs, MaSi, It, Ju, Al, Bu, RK, Gr, Cr, AE, Tu, An, LS, IJAnMa (nt)Gr, AE, An, LS, LJLiIJ, LS, ?An

CYCo, Sa, Me (R), Si, It, Gr, Cr, AE, An, Cy, LS, IJ, Li, Ga (E)An; only known from a few mountain peaks in west AnatoliaIt, Lu, Al, Gr, Cr, An, Br

Ma (R)Hs, Ma

CY (RIMaGa, Co, ItGa, It, Ju, (Hs?)Ag, Hs, Lu, Ma, TnAg (R), Co (probably introduced)AnAl, Bu, Ga, Gr, It, Ju, Si, TuLS, IJAn, Bu, RK, Tu.Lu, HsHsAnAn (Ex); probably identical with Rosularia sempervivum but type not yet traced

CY 6-NHs, Lu (R); species is confined to a specific biotope in north Portugal and a very small region

in adjacent SpainTh, Ag, MaAnall but Sn, Eg; extremely polymorphic species (cytologically as well as morphologically,

especially at the tetraploid level in the east Mediterranean region)Hs, Ga, Co, Sa, Si, It, JuAE, AnAn, IJ, LS; probably could be included in S. hispanicum, but taxonomic status not fully

understood; most likely related or conspecific with S. longibracteatumall but Bu, RK, Sn, Eg

168

R

RR

S. sempervivoidesS. sexangulareS. spuriumS. steudeliiS. stefcoS. stella turnS. stoloniferumS. subulatumS. surculosumS. tenellumS. tristria turnS. tuberiferumS. tu berosumS. tymphaeum

R

RR

RntR

RntRRR

R

RRntR

R

RR

R

R

3’. urstS. urvilleiS. versicolorS. villosumS. wilczekianumJovibarba allioniiJ. arenariaJ. heuflelliiJ. hirtaSempervivum arachnoideumS. armenumS. artvinenseS. ballsiS. brevipilumS. ciliosumS. davisiiS. dolomiticumS. erythraeumS. furseorumS. gillianiiS. gla brifoliumS. grandiflorumS. ispartaeS. italicumS. ja kucsiiS. kindingeriS. kosaniniiS. leucanthumS. macedonicumS. marmoreumS. minusS. montanumS. octopodesS. pisidicumS. ruthenicumS. staintoniiS. tectorumS. thompsonianumS. transcaucasicumS. vicentei ssp. cantabricumS. vicen tei ssp. pauiS. vicentei ssp. vicenteiS. wulfeniiUmbilicus albido-opacusU. chloranthisU. erectus

AnBu, Gr, Ga, It, Ju, AlAnAn, LSBu (R), GrBl, Ga, Co, Sa, Me, Si, It, Ju, Al, Gr, An (V), Ae (R)AnAnMaAnGr (R), CrBu, Gr

Tn, AgGr; confined to three mountain peaks in the Northern PindcAnJu, Al, Bu, RK, Gr, An, LS, IJMaHs, Ga, Co, Sa, It, Ju, Ag, Ma

Ma (RIGa (R); It(R)ItJu, Al, Bu, GrItHs, Ga, Co, It,AnAn

(3 wAnJu (R), Bu (R), Gr (R)AnHs, Bl, Ga, It (R)Al

An (R)An (R)An (RIIt

An wItAlJu (R), Gr (R)Ju (R)Bu

Ju CR)Ju, Al, Bu, Gr

An CR)Hs, Bl, Ga, It

Ju wAn (R)Ju, Bu, RK, Gr

An CR)Hs, Bl, Ga, It, Ju

Ju (RIAnHsHsHsItAEJu, Gr, AEIt, Ju, Al, Bu, Gr, Cr, Tu, An, LS

169

U. heylundiunusU. horizantulisU. gudituraus

Hs, Bl, Ma

u. in termediusU. mirus

all but Lu, Ga, Co, Gr, Tu, IJ, SnHSAn, LS, IJ, SnLi

U. neglectus Lu, Hs, BlU. purvijb-us Gr, Cr, AE, ANU. rupestris all but RK, IJ, EgU. tropuelifolius An, LS

EUPHORBIACEAEnt Euphorbiu ohtusijoliunt E. ojjkinurum ssp. ojjficinarumnt E. officinurum SSQ. echinusMa (nt)

MaMa

’ IUCN Red Data categories of global threa t arc given as recorded in the Plants Database maintained by WCMCare given by Focke Albers and IIenk VHart. Country categories are given in brackets following the country code.

as of July 1996. Additional C21 tegories

’ In compiling this list, all the succulents recorded by Grcuter, Burdet and Long (1984) in Med-Checklist were included, extended to the administrativelimits of the countries boarding the Mediterranean Sea, plus Portugal, Crimea, Bulgaria, and Jordan. The definition of succulent followed is that givenby Willert et al. (1992). The listing for Crassulaceae has been updated by Henk t’ Hart, and following Eggli (1988).

’ The two-letter distribution symbols are as used by Grcutcr, Burdet, and Long (1984):

AE East Aegean Islands, Ag Algeria, Al Albania, An Asiatic Turkey, Bl Balearic Islands, Bu Bulgaria, Co Corsica, Cr Crete and Karpathos, Cy Cyprus,Eg Egypt, Ga France, Gr Greece, Hs Spain, IJ Israel and Jordan, It Italy, Ju former Jugoslavia, Li Libya, LS Lebanon and Syria, Lu Portugal,Ma Morocco, Me Malta, RK Crimea, Sa Sardinia, Si Sicily, Sn Sinai, Tn Tunisia, Tu Turkey-in-Europe

Lampedusa Island,Italy.

170

Annex 6

Succulents of the Canary Islands

Prepared by David Bramwell.

Status Taxon Distribution

ntntnt

ntnt

ntE

Vnt

ntntnt

nt

ntE

CRASSULACEAEAeonium canarienseA. castello-paivaeA. cilia turn

A. cuneatumA. davidbram wellii

A. decorumA. gomerense

A. goochiaeA. ha worthii

A. hierrenseA. holochrysumA. luncerottense

A. lindleyi

A. manriqueorumA. mascaense

A. nobile

A. palmense

A. percarneumA. rubrolinea turn

E

nt

ntR

nt

ntnt

nt

nt

A. saundersii

A. sedifolium

A. simsiiA. smithii

A. sputhulatum

A. subplanumA. ta buliforme

A. undula turn

A. urbicum

nt A. valverdenseV A. vestitum

nt A. virgineum

Common on the north coast of Tenerife though declining.Very frequent on the northern slopes of La Gomera.Common in the Anaga region of Tenerife where hybridization with A. urbic‘um is the main threat. Its main

habitats in the forest regions (where it remains pure) are within the network of protcctcd arcas.Locally abundant in the laurel forests of Tenerife most of which arc now protected arcas.This is the most common Aeonium on La Palma and is extremely abundant cspccially in the sout 11 01’ the

island.Locally common on La Gomera, polymorphic.Extremely rare and endangered species, the main locality is threatened by a road-building project financed

by the EEC. It is endemic to a small area in the north-east of La Gomcra.Included here because of the recent fires which may have destroyed much of its natural habitat.Frequent in some areas of the north-west of Tenerife, many localities within the network of protcctcd arcas,

for example, Masca and Teno.Common on El Hierro.Common throughout the lower zone of Tenerife, La Palma, El Hierro and parts of La Gomcra.A species in expansion, for example, on recent lava flows. Most habitats protected and included in the MAB

reserve.Common in the lower zones of the Anaga region of Tenerife and also occurring on the west side of La

Palma.Very common all over Gran Canaria.Known from two populations in the Masca valley on Tenerife and thrcatencd by over-collecting. Probably

the rarest of all the Aeoniums but it is in cultivation and its habitat is included in a proposal for aprotected area.

Several populations exist, some of them quite large with over SW individuals but this spccics is thrcatcncdby overcollecting and by road improvement projects.

Common on La Palma but affected by recent forest fires, its status needs revising. The local form of thisspecies on El Hierro is common.

Locally very common throughout Gran Canaria.Rare with restricted distribution on La Gomera, threats include changes in agricultural methods and

overcollecting, some populations are protected by the National Park but the main ones arc outside thelimits and should be considered for reserve status especially Lomo de Carret6n and Bcnchijigua.

A locally very restricted species, abundant in a couple of small areas such as the Barranco dc la Laja whcrc itshould be strictly protected.

Abundant on the west side of Tenerife and parts of La Palma. Though of restricted distribution the spccicsis not threatened and most populations are in protected areas.

Common in the mountains of Gran Canaria.A local species of the southern parts of Tenerife from SW to 2000 m. Not common but most habitats are

included within protected areas.Found on all the western and central islands but common only on Tcnerife. Confined to a few I~xxlities on

La Gomera and Gran Canaria but all within the network of protected areas.Locally very common in the forest zones of La Gomera and widespread in the Gara.ionay National Park.Still frequent on the north coast of Tencrife but declining due to urbanization and general degradation ot

the natural habitat, not currently threatened.Locally abundant in the mountain regions of Gran Canaria and not thrcatcned cvcn though it is, ill some

areas, susceptible to hybridization with A. simsii.Common on the north and western slopes of Tenerife where its main threat is hybridization with 11. cilirrtrrrlr

in the north east, in other areas very abundant. The populations (rare) on La Gomcra may bc rcf’crablc toa distinct taxon and need further study.

Locally common on El Hierro.A species requiring revision and which may be only a local form of A. Iroloc~l~~~~.srrr~~ but if ;I good spccicx it is

rare, occurring only in the north east of La Palma where its main habitats may have been damaged bvrecent forest fires.

Locally frequent in the forest remnants of the island of Gran Canaria and extending to the Iowcr zone onthe west side, many of its main localities fall within protected areas.

171

ntnt

V

V

ntWV

RntR

nt

nt

Vntntnt

ntnt

KntVntntVKnt

VK

nt

nt

ntntE

V

ntnt

V

V

E

A. vista turnAichryson

bethencourtianumA. bollei

A. brevipetalum

A. laxumA. pachycaulon

A. palmensisA. parla toreiA. porphyrogennetos

A. puncta turn

A. tortuosumMonanthes

Monanthes adenoscepesM. amydrosM. anagensisM. brachycaulon

M. dasyphyllaM. laxiflora

M. minimaM. muralisM. niphophilaM. pallensM. polyphyllaM. praegeriM. purpurascensM. subcrassicaulis

M. wildpretiiSedum lancerottenseGreenovia

EUPHORBIACEAEEuphorbia aphylla

E. atropurpurea

E. balsamiferaE. berthelotiiE. bourgaeana

E. bravoana

E. broussonetiiE. canariensis

E. handiensis

E. lambii

E. mellqera

Locally frequent on the northern side of La Gomera.Restricted distribution on Fuerteventura but locally common.

Occurs on La Palma where recent forest fires have almost certainly affected all the main populations of thisspecies.

As previous species, this local La Palma endemic is seriously threatened by the effects of forest fires whichrecently destroyed almost 50% of the forests of the island.

Occurs on Tenerife, Gran Canaria, La Palma, La Gomera and El Hierro, very common.A rare species with individual endemic island subspecies which form part of the diversity of the species and

should be protected. The species is probably not monophyletic, each subspecies probably being ofindependent origin and so it is an important taxon from an evolutionary point of view.

Common on La Palma but declining along with its forest habitats.Occurs on Tenerife, Gran Canaria, La Palma, La Gomera and El Hierro, common and colonising walls.Sometimes confused with the previous species, seems to be in an expansion phase in the north of Gran

Canaria.Occurs on Tenerife, Gran Canaria, La Palma, La Gomera and El Hierro, very common but polymorphic and

its genetic diversity should be taken into account for conservation.Common cliff plant of Lanzarote and Fuerteventura.About 17 species but in need of a good revision; recent attempts have only added to the problems and the

basis provided by Praeger in the 1930s is still valid.A few scattered populations in the south of Tenerife, over-collected.Common on the north side of La Gomera.Still locally quite common but confined to the Anaga hills at the eastern end of Tenerife.Occurs on Tenerife and Gran Canaria, common but polymorphic and conservation measures should take

the possible genetic diversity into consideration.Tenerife, restricted distribution but locally very abundant.Occurs on all the islands; any conservation plan for this species should, however, take into account its

extreme polymorphism.May be the same as M. adenoscepes but needs revising.Occurs on Tenerife, La Palma and La Gomera, locally common.High mountain species, over-collected but now in Teide National Park and protected.Occurs on Tenerife, locally common.Occurs on Tenerife, very common.Single locality on the north coast of Tenerife.Occurs on Gran Canaria; this is probably one of the many local forms of M. brachycaulon.Occurs on Tenerife and La Gomera; needs taxonomic revision, appears to be common but confused with M.

muralis.Tenerife, single locality, recently described species.Locally abundant on the Famara cliffs of Lanzarote and not threatened. Main habitat is in a protected area.This genus of four species has no threatened taxa even though two species are restricted in distribution,

G. dodrentalis (its western Tenerife localities are in protected areas) and G. aizoon, protected in theGuimar natural park.

Tenerife, La Gomera, Gran Canaria, protected at Teno, Tenerife and the largest populations on GranCanaria will come within the proposed Roque Nublo National Park.

Abundant in the west and south of Tenerife where many populations are within the protected areasnetwork.

Widespread colonizing species abundant on all the islands in coastal regions.Endemic to La Gomera but locally common especially in the east and southeast.Remains, this species should be brought into cultivation and restocking used to augment the natural

population.Two main localities, Riscos de Agulo with few plants but in a protected area and Majona which is a Natural

Park with a relatively large population. A recovery plan should be included in the Park managementprogramme.

Tenerife, La Palma, La Gomera, El Hierro, the comment under the previous species also applies here.Common on most islands but rather rarer on La Gomera and with only a few localities on Lanzarotc

(Malpais de la Corona) and Fuerteventura (mainly Jandia).Declining species threatened by tourist development and over-collecting, its main remaining populations arc

protected in theory but are still declining especially due to the removal of young plants. The species shouldbe strictly protected in its natural habitat.

A local segregate of the previous species Gomera where three main populations occur; two of these arcwithin the National Park and if correctly managed this species should survive without too many problems.

Extremely rare species of Tenerife, La Palma and La Gomera. It is almost extinct in all its known localitieswhere populations are reduced to single figures with 1-5 individuals only. All localities should be protectedand reintroduction considered.

172

E

nt

E. regis-jubae

ASCLEPIADACEAECaralluma burchardii

Ceropegia cera tophora

c. chyantha

C. dichotomaC. fusca

C. hians

C. kra inziiCOMPOSITAE

Kleinia nerifolia(Senecio kleinia)

Occurs on Gran Canaria, Lanzarote, Fuerteventura; locally common but variable and a system of populationprotection for a wide sample of its diversity should be devised.

Confined to Lanzarote and Fuerteventura, most populations are in decline especially where rcgcncratingvegetation causes excessive shading. Its main localities should be protected and the species positivelymanaged.Endemic to La Gomera and confined to a few localities though further exploration of the southwest mayreveal new populations. Threatened by overcollection. Habitat should be protected and a management plandeveloped.Confined to a single locality in the south of Tenerife and was thought to be extinct. The remainingpopulation should be strictly protected and a recovery plan developed including ex situ conservation andrestocking.Tenerife, locally common especially on the north side, many populations in protected areas.Abundant in the south of Tenerife and on Gran Canaria but over-collected and sometimes taken from thewild for use in gardens, for example of hotels.A local segregate of the above from La Palma where it is locally common and El Hierro where it is confinedto the cliffs of El Golfo.North east part of La Gomera, habitats should be protected and a management plan developed.

A very common species on all the islands, it has a wide range of variability and a plan for the conservation ofits ample genetic diversity should be prepared.

173

Annex 7

Succulents of Madagascar

Compiled by Diedrich Supthut and Reto Nyffeler, October 1994.

An asterisk (*) indicates taxa found in trade. When a province is not indicated for infraspecific taxa, assume the same distribution as the species.Province abbreviations: Ant Antananarivo, Ants Antsiranana, Fia Fianarantsoa, Mah Mahajanga, Toa Toamasina, To1 Toliara, un unknown.


nt *R*E”RRnt *R”E”RK”R’”nt *KE*nt *R*nt *nt *RE’”E”E”R*E”R*K*KRK*nt *nt *E”nt *KKR*RE”KR’”R”EKR0nt *RR”RK

ALOACEAEAloe acutissima H.Perrier var. acutissima

- var. antanimorensis ReynoldsA. albiflora GuillauminA. alfiedii RauhA. andringitrensis H.PerrierA. antandroi (Decary) H.PerrierA. bakeri Scott-ElliotA. bellatula ReynoldsA. betsiliensis H.PerrierA. boiteaui GuillauminA. buchlohii RauhA. bulbillqera H.Perrier var. bulbillifera

- var. pauliana ReynoldsA. calcairophila ReynoldsA. capitata Baker var. capitata

- var. cipolinicola H.Perrier- var. gneissicola H.Perrier- var. quartziticola H.Perrier- var. silvicola H.Pcrricr

A. compressa H.Perricr var. compressa- var. rugosquamosa H.Perrier- var. schistophila H.Perrier

A. conifera H.PerrierA. cremersii LavranosA. cryptoflora ReynoldsA. decaryi GuillauminA. decorsei H. PerrierA. delphinensis RauhA. deltoideodonta Baker var. deltoideodonta

- var. breviflora H.Perrier- var. candicans H.Perrier

A. descoingsii ReynoldsA. divaricata A.Bergcr var. divaricata

- var. rosea (Decary) ReynoldsA. ericetorum BosserA. evthrophylla BosserA. jievetii ReynoldsA. fragilis Lavranos & RiiiisliA. guillaumetii CremersA. haworthioides Baker var. haworthioides

- var. aurantiaca H.PerrierA. helenae DanguyA. humbertii H.PerrierA. ibitiensis H.PerrierA. imalotensis ReynoldsA. isaloensis H.PerrierA. itremensis ReynoldsA. laeta A.Berger var. laeta

- var. maniaensis H.PerrierA. leandrii Bosser

Fia/TolTo1To1AntFiaTo1To1FiaTo1To1To1MahMahFiaAntFia

Mah/AntFia

MahAntAntAntFiaFiaFiaTo1FiaTo1FiaTo1FiaTo1

Fia/Tol/MahTo1MahFiaFia

AntsAntsFiaFiaTo1To1AntFia

Fia/TolFiaAntFiaToa

Distribution

K”KKKR’”E*E’”RKR*K*Knt *ntER”nt *KntR*

R*RRKRKRRRRR

A. macroclada BakerA. madecassa H.Perrier var. l?zacleczs.sa

- var. lutea GuillauminA. mayottensis A.BergerA. millotii ReynoldsA. parallelifolia H.PerrierA. parvula A.BergerA. perrieri ReynoldsA. peyrierasii CremersA. rauhii ReynoldsA. schomeri RauhA. silicicola H.PerrierA. suarezensis H.PerrierA. subacutissima G.D.RowleyA. suzannae DecaryA. trachyticola (H.Perrier) ReynoldsA. vaombe Decorse & Poiss. var. IYIOI?I~C’

- var. poissonii DecaryA. vaotsanda DecaryA.versicolor GuillauminA. viguieri H.PerrierLomatophyllum antsingyense LeandriL. belavenokense Rauh & R. GeroldL. citreum GuillauminL. occidentale H.PerrierL. oligophyllum (Baker) H.PerrierL. orientale H.PerrierL. prostratum H.PerrierL. viviparum H.PerrierL. roseum H.PerrierL. sociale H.Perrier

Ant/FiaAntu 11

ComoresTolAntFiaFia

AntsTo1TolAntAnts

Tol/FiaTel

Ant/FiaTo1To1TelTelTelMahTo1

Ll 1-l

MahLl I1

FiaMahAntsMahMah

All Lomatophyllum species grow at localities half-shaded by trcos and

shrubs. They are semi-succulents, have beautiful flowers, but are not wellknown by succulent enthusiasts.

APOCYNACEAEE* Pachypodium ambongense Poiss.R* P. baronii Constantin & Bois var. buroniiE” _ var. windsori (Poiss.) Pichonnt * P. brevicaule BakerE” P. decayi Poiss.nt * P. densiflorum Baker var. &~~.sijk)~-~~~~~R” - var. brevicalyx H.Perriernt * P. geayi Constantin & BoisE* P. inopinutum Lavranosnt * P. lamerei Drake var. lamereiR” _ var. ramosum (Constantin & Bois) PichonR” P. rosulatum Baker var. rosulatumnt * - var. delphinensis

(=stenanthum) Constantin & Bois

MahMahAnts

Ant/FiaMaI1

Ant/FiaAntTo1

Ma11Fia/Tol

TelMah

Tel

174

nt *nt *nt *nt *nt *nt *

- var. drukei (Constantin & Bois) Pichon- var. grucilius H.Perrier- var. horombense (Poiss.) G.D.Rowley

P. rutenbergianum Vatke var. rutenbergianum- var. meridionale (Pichon)H.Perrier

P. sofiense (Poiss.)H.Perrier

ASCLEPIADACEAEE*E”E”E”E”E”E

RRR*E”RRRRR”IsR”R*R”R*R*R”R”R*

Ceropegia armandii Rauh To1C. petignatii Rauh To1C. ho@uetteri Rauh MahC. simoneae Rauh To1C. bosseri Rauh & Buchloh FiaC. ruzajindratsirana (Rauh & Buchloh) Rauh FiaC. ampliatu var. madagascariensis Fia

Lavranos & MoratC. albisepta Jum. & H.PerrierC. viridis ChouxC. dimorpha RauhC. leroyi Rauh & Marn.-Lap.C. humbertii H.HuberC. scabra Jum. & H.PerrierC. saxatilis Jum. & H.PerrierC. madagascariensis Decne.Stapelianthus arenarius Bosser & MoratS. calcarophilus MoratS. decavi ChouxS. hardyi LavranosS. insignis Desc. var. insignis

- var. tangoboryensis RauhS. keraudreniae Bosser & MoratS. mudagascariensis (Choux) ChouxS. montagnacii (Boiteau) Boiteau & BertrandS. pilosus Lavranos & Hardy

To1To1FiaFia

AntsTo1

MahMahTo1

?To1To1To1To1To1To1To1To1

MahFia/TolFia/Tol

Mah/AntsTo1Mah

Most Stapelianthus species grow in the Euphorbia-Didieraceen-forest inthe south-west of Madagascar, which is threatened by clearing and fire.Stapelianthus species are not easy to grow, all are succulent.

Succulent species (following S. Liede 1992): low-growing and leaflessR Cynanchum descoingsii Rau h To1E” C. rossii Rauh To1K C. antandroy Desc. To1R* C. compactum Choux FiaK C. juliani-murnieri Desc. To1R” C. macrolobum Jum. & H.Perrier To1R* C. marnieranum Rauh To1K* C. perrieri Choux FiaR C. rauhianum Desc. FiaK Folotsia sarcostemmoides Cost. & Bois To1E Karimbolea verrucosa Desc. To1K Sarcostemma insignis N.E.Br. AntSucculent species: leafless vinesK Cynanchum aequilongum Choux Tol/ToaK C. ambositrense Choux FiaK C. ampanihense Jum. & H.Perrier To1K C. arenarium Jum. & H.Perrier MahK C. bekinolense Choux FiaK C. bisinuatum Jum. & H.Perrier To1K C. decaisneanum Desc. Fia/Tol/MahK C. gerrerdii (Harvey)Liede To1K C. implicatum (Jum. & H.Perrier) Ants

Jum. & H.PerrierK C. lecontei Choux To1

K C. luteifluens (Jum. & H.Perrier) Desc.K C. macranthum Jum. & H.PerrierK C. madecassum Desc.K C. mahafalense Jum. & H.PerrierK C. menarandrense Jum. & H.PerrierK C. messeri (Buchenau) Jum. & H.PcrrierK C. nodosum (Jum. & H.Perrier) Desc.K C. radiatum Jum. & H.PerrierK Folotsia aculeatum Jum. 81 H.PerrierK F. floribundum Desc.K F. grandiflurum Jum. & H.PerrierK F. madagascariense (Jum. & H.Perrier)DtK Sarcostemma decorsei Cost. & Gall.K S. madagascariensis Desc.K S. viminale R.Br.Herbaceous scramblers with tuberous rootsK Cynanchum subtilis LiedeK C. andringitrense ChouxK C. danguyanum ChouxK C. madagascariense K.Schum.K C. angkavokeliense ChouxK C. bojerianum Decne.K C. cucullatum N.E.Br.K C. jumellei ChouxK C. lineare N.E.Br.K C. moramangense ChouxK C. papilla turn C houxK C. surprisum LiedeK C. napiforme ChouxErect herbs with tuberous rootsK Cynanchum appendiculatum ChouxK C. fibriatum ChouxK C. napferum ChouxK C. junciforme (Decne.)LiedeK C. sessilijlorum (Decne.)LiedeBroad leaved vinesR C. pachylobum Choux

BALSAMINACEAER” Impatiens tuberosa H.Perrier

:sc.

TolTo1To1

Tol/FiaTo1

Ants/To1TolToITo1Fia

MahFia/Tol

To1To1Tol

To1Ant

Fia/TolAnt/Fia

AntAnt/Ants

AntAntAntAnt

Ant/FiaTo1Ll 11

FiaAntAnt

Ant/Fia/ToIAnt/Mall

TO I

Ants

COMPOSITAE (Asteraceac) (Group XVI ex Fl. dc Madagascar‘)K”KKKR”KR”K*KKKK”

KKKKR*KKKK

Senecio antandroi Scott-Elliot TolS. antitensis Baker FiaS. barorum Humbcrt Fi a/To1S. boiteaui Humbert TolS. canaliculatus DC. Ant/FiaS. capuronii Humbert FiaS. cedrorum Raynal TolS. crassissimus Humbert Fia/TolS. decuryi Humbert FiaS. hirto-crassus Humbert Ant/FiaS. hildebrandtii Baker AntiFiaS. longiflorus (DC.) Schultz-Bip.

- var. madugascariensis (Humbert) G.D.Rowlcy To1S. marnieri Humbert AntS. melastomaefolius Baker ssp. rllt’lrrstol?lrrl’fi~lillS AntJFia

- ssp. longibracteatus Humbert AntsS. mesenzb~a?lt~lenloines Bojer ex DC. Ant/F&S. meuselii Rauh FiaS. navicularis Humbert TolS.quartzitico1u.s Humbcrt Ant/FiaS. saboureaui Humbert AnfsS. sakamaliensis (Humbert)Humbert To1

175


K S. sakalavorum HumbertK” Kleinia descoingii (Humbert)JeffreyK” K. madagascariensis (Humbert) P.HallidayK Notonia hehdingii Rauh & Buchloh

CRASSl JLACEAE

MahFia/Tol

Mah/TolTo1

RKR”KKKK

K

KK”KK’”KKnt *K”K*KKnt *K*KKKK

KK

nt *KR**

K*KK”KKK”K*Knt *KR’”*

K*K

K*KK

Crassula cordi$olia Baker AntC. fvagilis Baker unC. humbertii Desc. To1C. micans Vahl ex Baillon unC. nummulariifolia Baker AntKalanchoe adelae Raym.-Hamet ComoresK. adolphi-engleri Raym.-Hamet Fia/TolK. aliciae Raym.-Hamet see

K. pubescens BakerK. ambolensis Humbert To1K. amplexicaulis (Baker) Baillon

see K. campanulata (Baker) BaillonK. antanosiana Drake To1K. arborescens Humbert To1K. aromatica H.Perrier AntK. beauverdii Raym.-Hamet var. beauverdii Ant/Fia

- var. guignardii Raym.-Hamet Mahajanga- var. parviflora Manning & Boit. To1

K. beharensis Drake var. beharensis Tol/Fia- var. aureo-aeneus H.Jacobsen To1- var. subnuda H.Jacobsen To1

K. bergeri Raym.-Hamet var. bergeri Ant/Toa/Tol/Fia- var. glabra Manning & Boit.

K. bitteri Raym.-Hamet Ant/FiaK. blossfeldiana Poelln. AntsK. boissii Raym.-Hamet & H.Perrier Mah/TolK. bouvetii Raym.-Hamet & H.Perrier Mah/ToaK. bracteata Scott-Elliott To1K. campanulata (Baker) Baillon Ant/Fia

- var. campanulata- var. orthosyla Manning & Boit. Ant

K. chapototi Raym.-Hamet & H.Perrier MahK. constantinii Raym.-Hamet

see K. beauverdii Raym.-HametK. daigremontiana Raym.-Hamet & H.Perrier To1K. ebracteata Scott Elliott To1K. eriophylla Hils. & Bojer Fia/AntK. fedtschenkoi Raym.-Hamet & H.Perrier

- var. fedtschenkoi Fia/Ant/Tol- var. isalensis Manning & Boit.

K. gastonis-bonnieri Raym.-Hamet & H.PerrierK. gentyi Raym.-Hamet & H.Perrier Ant/FiaK. globulifera H.Perrier Ants/MahK. gracilipes (Baker)Baillon Tol/Fia/AntK. grandidieri Baillon To1K. heckelii Raym.-Hamet & H.Perrier AntK. hildebrandtii Baillon To1K. humberti H.Perrier To1K. integrifolia Baker Fia/AntK. jongmansii Raym.-Hamet & H.Perrier

- var. jongmansii Fia- var. ivohibensis Humbert

K. julea Raym.-Hamet & H.Perriersee K. beauverdii Raym.-Hamet

K. laxiflora Baker var. laxifiora- var. stipitata Manning & Boit.- var. subpeltata Manning & Boit

Ant/Fia

KR’”KK”K”KK”KKKKKKKK”K*

K”KK

KK

KKKKKKKKKKKK*K*KK”KKK

K*KKKnt *R”Knt *K

Knt *K*KKRK

- var. violacea Manning & Boit.K. linearifolia DrakeK. macrochlamys H.PerrierK. manginii Raym.-Hamet & H.PerrierK. marnierana H.JacobsenK. millotii Raym.-Hamet & H.PerrierK. miniata Hils. & Bojer var. miniata

- var. andringitrensis H.Perrier- var. anjirensis H.Perrier

TolAnts

An t/FL‘7

To1Ant/Fia/Tol

- var. comfertifolia H.Perrier- var. peltata Baker- var. sicaformis Manning & Boit.- var. subsessilis H.Perrier

K. mortagei Raym.-Hamet & H.PerrierK. nadyae Raym.-HametK. orgyalis BakerK. panduriformis (Baker) Baillon

see K. companulata (Baker) BaillonK. peltata (Baker)Baillon var. peltata

- var. mandrakensi H.Perrier- var. stapfii (H.Perrier) Raym.-Hamet

& H.PerrierK. poincarei Raym.-HametK. porphyrocalyx (Baker) Baillon

var. porphyrocalyx

AntAntsTol

Tol/Fia

Ant/Fia/MahAnt/Mah

Ants

Fia/TolAnt/Tea

- var. sambiramensis Humbert Ants- var. sulphurea Baker TOI

K. proZt$era (Bowie)Raym.-Hamet ToaK. pseudocampanulata Manning & Boit. FiaK. pubescens Baker var. pubescens Ant/Tol/Fia/Ants

- var. alexiana Manning & Boit. Fia- var. brevicazyx Manning & Boit.- var. decolorata Manning & Boit.- var. grandiflora Manning & Boit.- var. subglabrata Manning & Boit. To1- var. subsessilis Manning & Boit.

K. pumila Baker Ant/FiaK. rhombopilosa Manning & Boit. To1K. rolandi-bonapartei Raym.-Hamet & H.Perrier AntsK. rosei Raym.-Hamet & H.Perrier var. rosei Tol

- var. seyrigii Manning & Boit. Tol- var. vartfolia Guill. & Humbert Tol

K. rubella (Baker)Raym.-Hamet un

K. scandens H.Perriersee K. beauverdii Raym.-Hamet

K. schizophyllu (Baker) Baillon An t/FiaK. serrata Manning & Boit. FiaK. streptantha Baker Ants/MallK. suarezensis H.Perrier AntsK. synsepala Baker Fia/Ant/TolK. tetraphylla H.Perrier Fia/An tK. tieghemi Raym.-Hamct Ull

K. tomentosa Baker Fia/Ant/Tol/AntsK. trichantha Baker u I1

K. tsaratananensis H.Perriersee K. rolandi-bonaparteiRaym.-Hamet & H.Pcrrier

K. tuberosa H.Perrier AntK. tubiflora (Harvey)Raym.-Hamet Tol/FiaK. uniflora (Stapf)Raym.-Hamet var. IIII$OW Ants

- var. brachycaZyx Manning & Boit.K. verticillata Scott-Elliot To1K. viguieri Raym.-Hamet & H.Perrier To/MallK. wazdheimii Raym.-Hamet & H.Perrier FiaiTol

176


K Sedum madagascariense H.Perrier Fia

CUCURBITACEAEK Seyrigia bosseri Keraudren To1K 5’. gracilis Keraudren To1K S. humbertii Keraudren To1K S. multiflora Keraudren To1The clearing of the dry thorn-forest is a possible reason for thethreatening of the species of Seyrigia. They are not often found in trade.

nt * Trochomeriopsis diversifolia Ants/Mah/Tolnt Xerosicyos danguyi Humbert To1nt XI perrieri Humbert Mah/Tolnt X decalyi Guill. & Keraudren Mah/TolR* X pubescens Keraudren To1

(caudiciform, traded in large quantities)K” Zygosicyos tripartitus Humbert To1K 2. hirtellus Humbert To1Trochomeriopsis and Zygosicyos are in trade from time to time.

DIDIEREACEAEE* Alluaudia ascendens (Drake) Drakent” A. comosa Drakent A. dumosa Drakent A. humbertii ChouxR” A. montagnacii Rauhnt* A. procera DrakeR Alluaudiopsisfiherensis Humbert & Chouxnt A. marnierana Rauhnt” Decaryia madagascariensis Chouxnt” Didierea madagascariensis Baillonnt* D. trollii Capuron & Rauh

To1To1To1

Tol/FiaTo1To1To1To1To1To1To1

DRACAENACEAER Sansevieria canaliculata Carri&-eR S. sambiranensis H.Perrier

Sansevieria species are semi succulent.

AntsAnts

EUPHORBIACEAEKR*nt*R

Euphorbia alcicornis BakerE. alfiedii RauhE. alluaudii Drake ssp. alluaudii

- ssp. oncoclada (Drake) Friedm. & CremersE. ambatojinandranae Leandri

see E. stenoclada Baillon ssp.ambatofinandranae (Leandri) Cremers

E’” E. ambovombensis Rauh & Razaf.K E. anahaka Humbert & LeandriK E. analalavensis LeandriR E. annamarieae Rauhnt* E. ankarensis Boit.K E. arahaka Humbert & LeandriK” E. beharensis LeandriK E. benoisti LeandriK E. betacea HumbertK E. biaculeata DenisK E. boinensis Humbert & LeandriK E. boissieri BaillonK E. boiteaui LeandriK E. boivinsis Humbert & LeandriR’” E. bongola vensis Rau hK” E. bosseri LeandriK E. brachyphylla DenisR* E. bulbispina Rauh & Razaf.

unAntsTo1To1

To1un

AntsTo1

AntsTo1To1ununTo1MahunTo1unTo1To1To1

Ants


nt *KKE’”Ent*E”

E”E*E””E*EE*K”KntI”nt *Knt *nt*Knt

ntKnt *nt*E”KKKnt*R”R*Knt*R”R*nt *Knt

K*R’”KKK

K”KK”ntKKKntK

E. capsaintemariensis Rauhsee E. decaryi Guillauminvar. capsaintemariensis (Rauh) Cremers

E. capuronii Ursch & LeandriE. caputaureum DenisE. commersonii DenisE. cremersii Rauh & Razaf.

-fa. viridifolia RauhE. croizatii LeandriE. cylindrifolia Marn.-Lap. & Rauh

- ssp. cylindrif~~lia- ssp. tuberifera Rauh

E. decaryi Guillaumin var. decatyi- var. ampanihensis Cremers- var. capsaintemariensis (Rauh) Cremers- var. robinsonii Cremers- var. spirosticha Rauh & Buchloh

E. delphinensis Ursch 81 LeandriE. denisiana GuillauminE. denisii OudejansE. didiereoides LeandriE. duranii Ursch & Leandri var. duranii

- var. ankaratrae Ursch & LeandriE. enterophora Drake ssp. enterophora

- ssp. crassa CremersE. eumyrodes BaronE. famatamboay Friedm. & Cremers

- ssp. famatamboay- ssp. itampolensis Friedm. & Cremers

E. fauroti Franch.E. jianarantsoae Ursch & LeandriE. fiherenensis Poiss.E. francoisii Lcandri var. j?ancoisiiE. j?-ancoisii var. rakotoza-fii CremersE. geayi Const. & Gall.E. genistoides Berg.E. genoudiana Ursch 8r LeandriE. gottlebei RauhE. guillauminiana Boit.E. guillemetii Ursch & LeandriE. hedyotoides N.E.Br.E. herman-sch wartzii RauhE. hofstaetteri RauhE. horombensis Ursch Rc LcandriE. imerina CremersE. intisy Drake var. intisy

- var. maintyi (Decorse) Poiss.see E. maintyi Leandri

E. isaloensis DrakeE. kondoi Rauh & Razaf.E. laro DrakeE. leandriana Boit.E. leptomyuru Poiss.E. leucodendron Drake

see E. ulluaudii Drake ssp. allrraudiiE. leuconeuru Boiss.E. lohahensis Grand.E. lophogona Lam. var. lophogona

- var. tenuicaulis RauhE. macroglypha LemaireE. mahafalcnsis Leandri var. I?lllllc!~~l~~rlsi.s

- var. xanthadenia (Denis)LeandriE. maintyi LeandriE. mandrariensis Drake

TolToITolTo1To1To1To1Ll 1-l

TolFiaFiaAntTO1FiaI1 11

TolTol

11 I-l

FiaTolTO1Ll I1

Ll I1

1111

TolTo1MallTolTo1AntsTol

Fia/TolTO ITol

Fia/TolTolu 11

‘)

11 I7

L1 I1

1111

TO ITol1111

TolTo1TolTo1

177

Distribution

KKKK*KKKK”KK”KKK*K*R”R”K”

E”EEER”nt *R”

K*KR”E*nt *nt”R’”R*nt *R”R”0”0KKE”KKR*KKR”KKnt *R”KKKKnt*KE”nt”

E. mandraviob LeandriE. mangokyensis DenisE. melanacantha DrakeE. milii Des Moul. var. milii

unTo1un

Madagascar- var. betsiliana Leandri- var. bevilaniensis (Croizat) Ursch & Leandri- var. bosseri Rauh syn. see E. neobosseri Rauh- var. breonii (Nois.) Ursch & Leandri- var. hislopii (N.E.Br.)- var. imperatae (Leandri) Ursch & Leandri- var. longifolia Rauh- var. roseana Marn.-Lap.- var. splendens (Hooker) Ursch-Leandri- var. tananarivae Leandri- var. tenuispina Rauh & Razaf.- var. tulearensis Ursch & Leandri- var. vulcanii Leandri

E. millotii Ursch & LeandriE. moratti Rauh var. moratti

- var. antsingiensis Cremers- var. bemarahensis Cremers- var. multiflora Rauh

E. neobosseri RauhE. neohumbertii Boit. var. neohumbertii

- var. aureo-viridiflora RauhE. obcordata Denis (later homonym)

see E. denisii OudejansE. oncoclada Drake see

E. alluaudii Drake ssp. oncoclada(Drake) Friedm. & Cremers

E. orthoclada Baker ssp. orthoclada- ssp. vepretorum (Drake) Leandri

E. pachypodioides Boi t.E. parvicya tophora Rau hE. pauliana Ursch & LeandriE. pedilanthoides DenisE. perrieri Drake var. perrieri

- var. elongata DenisE. plagiantha DrakeE. platyclada Rauh var. platyclada

- var. hardyi RauhE. primulifolia Baker var. primulifolia

- var. begardii CremersE. prostrata PoissE. pyrifolia Lam k.E. quartziticola LeandriE. radula Poiss.E. razafinjohanii Ursch & LeandriE. rossii Rauh & BuchlohE. rubrostriata Drake var. rubrostriata

- var. dulioti DrakeE. sakarahaensis RauhE. sapiifolia Poiss.E. solota LeandriE. stenoclada Baillon ssp. stenoclada

FiaTo1R*

AntsMahMahMahMahTo1

AntsAnts

Fia/AntTo1

AntsTo1MahMahMahAntsTo1To1To1

Ant/FiaFiaun

AntsFiaunFiaTo1ununTo1ununFiaP.- ssp. ambatofinandranae (Leandri) Cremers ria

E. suareziana Croizat AntsE. tardieuana Leandri ToaE. tetraptera Poiss. unE. thymifolia Poiss. unE. tirucalli L. To1E. tsimbazatae Leandri unE. tulearensis (Rau h) Rau h To1E. viguieri Denis var. viguieri Ants/Mah


R” - var. ankarafantsiensis Ursch & Leandri Mahnt* - var. capuroniana Ursch & Leandri AntsK - var. tsimbazazae Ursch & Leandri unR” - var. vilanandrensis Ursch & Leandri Mah

E. xylophylloides Lemairesee E. enterophora Drake var. crassa Cremers

K E. zakamenae Leandri Toa

PASSIFLORACEAEK Adenia ambongensis Claverie Mah/TolK A. antongilliana Schinz Ant s/TeaK” A. epigea H.Perrier MahK A. elegans H.Perrier To1K A. densiflora Harms An t/FiaK” A. firingalavensis (Drake)Harms Antsilanana/Mah/TolK A. monadelpha H.Perrier To1K* A. olaboensis Claverie Ants/Mah/TolK* A. peltata Schinz Tol/FiaK” A. perrieri Claverie MahK A. refracta Schinz MallK A. subsessilifolia H.Perrier To1K A. sphaerocarpa Claverie Mah/TolK - ssp. mandrarensis H.Perrier To1K - ssp. isaloensis H.Perrier Fia/TolAll Adenia species are caudiciform succulents, which arc in trade as bigand adult specimens.

PEDALIACEAEnt * Uncarina abbreviata (Baillon) Ihlenf. & Straka Tol/Mahnt” U. decaryii Humbertnt* U. grandidieri (Baillon) Stapfnt U leandrii Humbertnt U. leptoccarpa (Decne.) Ihlenf. & Strakant * U. peltata (Baker) Stapfnt U. perrieri HumbertE* U. roeoesliana Rauhnt U. sakalava Humbertnt * U. stellulifera Humbert

VITACEAERKKKKKKR*KKKnt”nt”KKKntK*KKKKK

Cyphostemma pachypus Desc.C. ankirihitrense Desc.C. amplexicaule Desc.C. comigerum Desc.C. cristigerum Desc.C. delphinense Desc.C. echinocarpum Desc.C. elephantopus Desc.C. glandulosopilosum Desc.C. greveanum Desc.C. horombense Desc.C. laza Desc. var. laza

- var. parvifolia Desc.C. leucorufescens Desc.C. macrocarpum Desc.C. manambovense Desc.C. microdipterum Desc.C. montagnacii Desc.C. pumilum Desc.C. roseiglandulosa Desc.C. sakalavense Desc.C. tsaratananense Desc.C. vezense Desc.

To1Tol

Fia/TolMah/TolAnt/MahAnt/Mah

MallTo1

AntsMahAntMahFiaTolTolTo1

Ants/TeaAnts/Mah/Fia

FiaTo1TolAntAntsTol

Tol/Ant/FiaTo1To1MahToIAntsTo1

178

Annex 8

Threatened succulents recorded for the Floraof southern Africa (FSA) region

Compiled by Craig Hilton-Taylor.

The data presented here were extracted from the threatened plantsdatabase ‘SARARES’ maintained at the Ecology and Conservationresearch division of the National Botanical Institute in Kirstenbosch,South Africa. All species are endemic unless noted by an asterisk (*)<

AIZOACEAERRRRRKKKKRKIKKKIRRRRVExRR/VKRVKKKIRVVIVIExRRVRRRRVRRRR

Aloinopsis acuta L.BolusA. jamesii L.BolusA. loganii L.BolusA. setifera (L.Bolus) L.BolusA. villetii (L.Bolus) L.BolusAqyroderma framesii L.Bolus ssp. fiamesiiA. ringens L.BolusA. subalbum (N.E.Br.) N.E.Br.A. testicular-e (Aiton) N.E.Br.Astridia citrina (L.Bolus) L.BolusA. herrei L.BolusA. speciosa L.BolusA. vanheerdei L.BolusBergeranthus addoensis L.BolusB. katbergensis L.BolusBijlia tugwelliae (L.Bolus) L.BolusBraunsia stayneri (L.Bolus) L.BolusB. vanrensburgii (L.Bolus) L.BolusCaryotophora skiatophytoides LeistnerCephalophyllum confusum (Dinter) Dinter & SchwantesC. jidleri L.BolusC. parvulum (Schltr.) H.E.K.HartmannC. pulchellum L.BolusC. pulchrum L.BolusC. rostellum (L.Bolus) H.E.K.HartmannC. slaminodiosum L.BolusC. tetrastichum H.E.K.HartmannChasmatophyllum braunsii SchwantesC. maninum L.BolusC. willowmorense L.BolusCheiridopsis alata L.BolusC. delphinoides S.A.HammerC. pearsonii N.E.Br.C. peculiaris N.E.Br.C. rudis L.BolusC. umdausensis L.BolusC. velox S.A.HammerCircandra serrata (L.) N.E.Br.Cleretum lyratifolium Ihlenf. & StruckConophytum achabense S.A.HammerC. acutum L.BolusC. armianum S.A.HammerC. auriflorum Tischer ssp. auriflorumC. bicarinatum L.BolusC. blandum L.BolusC. burgeri L.BolusC. carpianum L.BolusC. concavum L.BolusC. ernstii S.A.Hammer ssp. emstiiC. frutescens Schwantes

179

RERRKRRKRRExRRRExERRRV

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IVRREKERRRKRKIERIRRV

C. halenbergense (Dinter & Schwantes) N.E.Br.C. herreanthus S.A.Hammer ssp. hel7-eanthrrsC. khamiesbergense (L.Bolus) SchwantesC. klinghardtense Rawe ssp. baradii (Rawe) S.A.HammerC. lithopsoides L.Bolus ssp. lithopsoidesC. loeschianum TischerC. phoeniceum S.A.HammerC. praesectum N.E. Br.C. regale LavisC. ricardianum Loesch & Tischer ssp. ricardianlimC. ricardianum Loesch & Tischer ssp. rubrifkorum TischerC. rugosum S.A.Hammer ssp. rugosur?lC. rugosum S.A.Hammer ssp. sanguincrrm S.A.HammerC. schlechteri SchwantesC. semivestitum L.BolusC. smorenskaduense de Boer ssp. 1~mnariun~ S.A.HammcrC. smorenskaduense de Boer ssp. snzorcrzsk(rclrrcrlSC’C. swanepoelianum Rawe ssp. swane~~oelia)lrrr71C. taylorianum (Dinter & Schwantes) N.E.Br. ssp. taylorianumC. uviforme (Haw.) N.E.Br. ssp. srrbincanum (Tischer)

S.A.HammerC. vanheerdei TischerC. velutinum Schwantes ssp. velutinumC. verrucosum (Lavis) G.D. RowleyDelosperma clavipes LavisD. guthriei LavisD. hallii L.BolusD. inaequale L.BolusD. leendertziae N.E.Br.D. macrostigma L.BolusD. mariae L.BolusD. pondoense L.BolusD. subpetiolatum L.BolusD. suttoniae LavisD. velutinum L.BolusDidymaotus lapidij&mis (Marloth) N.E.Br.Dinteranthus microspermus (Dinter & Derenb.) Schwantcs ssp.

microspermusD. pole-evansii (N.E.Br.) SchwantcsD. vanzylii (L.Bolus) SchwantesD. wilmotianus L.Bolus ssp. impunctatus N.SaucrDiplosoma luckhofii (L.Bolus) Schwantes cx Ihlcnf.D. retroversum (Kensit) SchwantesDorotheanthus apetalus (L.f.) N.E.Br.Drosanthemum austricolum L.BolusD. bellum L.BolusD. hallii L.BolusD. micans (L.) SchwantesD. nordenstamii L.BolusD. thudichumii L.Bolus var. gracilis L.BolusEbracteola montis-moltkei (Dinter) Dinter C! SchwantcsEctotropis alpina N.E.Br.Erepsia brevipetala L.BolusE. dubia LiedeE. hallii L.BolusE. insignis (Schltr.) SchwantcsE. patula (Haw.) SchwantesE. pentagona (L.Bolus) L.Bolus

IERExIVVR

RKKExKKR/VKRRRWVRIRRVRK

R

R

IVERKKKKIVVRRIERRK

RKKK

R

K

K

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E. pillansii (Kensit) LiedeE. polita (L.Bolus) L.BolusE. poZypetala (A.Berger & Schltr.) L.BolusE. promontorii L.BolusE. steytlerae L.BolusE. villiersii L.BolusFaucaria tigrina (Haw.) SchwantesFenestraria rhopalophylla (Schltr. & Diels) N.E.Br. ssp.

aurantiaca (N.E.Br.) H.E.K.HartmannFrithia pulchra N.E.Br. var. pulchraGibbaeum angulipes (L.Bolus) N.E.Br.G. dispar N.E.Br.G. esterhuyseniae L.BolusG. nebrownii TischerG. pachypodium (Kensit) L.BolusG. schwantesii TischerImitaria rnuirii N.E.Br.Jacobsenia hallii L.BolusJensenobotrya lossowiana HerreJordaaniella clavifolia (L.Bolus) H.E.K.HartmannJ. uniflora (L.Bolus) H.E.K.HartmannLampranthus algoensis L.BolusL. dunensis (Sond.) L.BolusL. fugitans L.BolusL. rustii (A.Berger) N.E.Br.L. scaber (L.) N.E.Br.L. stenus (Haw.) N.E.Br.Lithops aucampiae L.Bolus ssp. aucampiae var. koelmanii (de

Boer) D.T.ColeL. aucampiae L.Bolus ssp. euniciae (de Boer) D.T.Cole var.

euniciaeL. aucampiae L.Bolus ssp. euniciae (de Boer) D.T.Cole var.

fluminalis D.T.ColeL. bromfieldii L.Bolus var. glaudinae (de Boer) D.T.ColeL. coleorum S.A.Hammer & R.UijsL. comptonii L.Bolus var. comptoniiL. comptonii L.Bolus var. weberi (L.Bolus) B.FearnL. dinteri Schwantes ssp. dinteri var. brevis (L.Bolus) B.FearnL. dinteri Schwantes ssp. dinteri var. dinteriL. dinteri Schwantes ssp.fiederici (D.T.Cole) D.T.ColeL. dinteri Schwantes ssp.multipunctata (de Boer) D.T.ColeL. divergens L.Bolus var. amethystina de BoerL. divergens L.Bolus var. divergensL. dorotheae NelL. franciscii (Dinter & Schwantes) N.E.Br.L. fillviceps (N.E.Br.) N.E.Br. var. lactinea D.T.ColeL. gesineae de Boer var. annae (de Boer) D.T.ColeL. gesineae de Boer var. gesineaeL. geyeri NelL. gracidelineata Dinter ssp. brandbergensis (de Boer) D.T.ColeL. gracilidelineata Dinter ssp. gracilidelineata var. waldroniae de

BoerL. helmutii L.BolusL. hookeri (A.Berger) Schwantes var. dabneri L.BolusL. hookeri (A.Berger) Schwantes var. marginata (Nel) D.T.ColeL. hookeri (A.Berger) Schwantes var. subfenestrata (de Boer)

D.T.ColeL. hookeri (A.Berger) Schwantes var. susannae (D.T.Cole)

D.T.ColeL. karasmontana (Dinter & Schwantes) N.E.Br. ssp.

karasmontana var. aiaisensis (de Boer) D.T.ColeL. karasmontana (Dinter & Schwantes) N.E.Br. ssp.

karasmontana var. tischeri D.T.ColeL. lesliei (N.E.Br.) N.E.Br. ssp. burchellii D.T.ColeL. lesliei (N.E.Br.) N.E.Br. ssp. lesliei var. hornii de BoerL. lesliei (N.E.Br.) N.E.Br. ssp. lesliei var. mariae D.T.Cole

R

R

VR

RRKRR/VREKKIRVRIKRER

R/VEIIIRIRKEEIER/VVIRKIIRRRIRRIIRIIREx

L. meyeri L.BolusL. naureeniae D.T.ColeL. olivacea L.Bolus var. nebrownii D.T.ColcL. otzeniana NelL. pseudotruncatella (A.Berger) N.E.Br. ssp. ~~sel~ciotn~~lc.Lrtell~~

var. elisabethae (Dinter) de Boer Ore BoomL. pseudotruncatella (A.Berger) N.E.Br. ssp. pscl~ciotrl~~~cat~~llrl

var. riehmerae D.T.ColeL. pseudotruncatella (A.Berger) N.E. Br. ssp. volkii (Schwantes

ex de Boer & Boom) D.T. ColeL. salicola N.E.Br.L. schwantesii Dinter ssp. schwantesii var. l-ugosa (Dinter) de

Boer & BoomL. viridis C.A.Liickh.L. werneri Schwantes & H. JacobsenMestoklema albanicum N.E.Br. ex GlenMitrophyllum abbreviatum L.Bolus44 roseum L.BolusMossia intervallaris (L.Bolus) N.E.Br.Muiria hortenseae N.E.Br.Nananthus pole-evansii N.E.Br.N. vittatus (N.E.Br.) SchwantesNelia pillansii (N.E.Br.) SchwantesN. schlechteri SchwantesNeohenricia sibbettii (L.Bolus) L.BolusN. spiculata S.A.HammerOdontophorus marlothii N.E.Br.Ophthalmophyllum longitubum L.Bolus0. villetii L.BolusOrthopterum coegana L.BolusPleiospilos compactus (Aiton) Schwantes ssp. minor (L.Bolus)

H.E.K.Hartman & LiedeP. nelii SchwantesP. simulans (Marloth) N.E.Br.Rabiea jamesii (L.Bolus) L.BolusRhinephyllum inaequale L.Bolus var. inaequaleR. inaequale L.Bolus var. latipetalum L.BolusRuschia amicorum (L.Bolus) SchwantesR. filamentosa (L.) L.BolusR. firma L.BolusR. lawsonii (L.Bolus) L.BolusR. leipoldtii L.BolusR. promontorii L.BolusR. rubricaulis (Haw.) L.BolusSaphesiaflaccida (Jacq.) N.E.Br.Schwantesia acutipetala L.BolusS. borcherdsii L.BolusS. triebneri L.BolusScopologena veruculata (L.) L.BolusStayneria neilii (L.Bolus) L.BolusStomatium geoffreyi L.BolusS. ronaldii L.BolusTanquana archeri (L.Bolus) H.E.K.Hartmann 81 LiedeT. hilmarii (L.Bolus) H.E.K.Hartmann & LiedeTrianthema hereroensis SchinzTrichodiadema aureum L.BolusT. burgeri L.BolusT. hallii L.BolusT. obliquum L.BolusT. peersii L.BolusT. pygmaeum L.BolusT. rogersiae L.BolusT. rupicolum L.BolusK!okia ater S.A.HammerZeuktophyllum suppositum (L.Bolus) N.E.Br.

180

ALOACEAEVVERERKRRRRRRVKRRRKRRKVVRRKRRVRRREEKRRVIRVVVRRR

IVRRRRRRKERVRRK

R

Aloe alhida (Stapf) ReynoldsA. arenicola ReynoldsA. bowiea Schult. & Schu1t.f.A. buhrii LavranosA. chlorantha LavranosA. comosa Marloth & A.BergerA. cooperi Baker ssp. pulchra Glen & D.S.HardyA. corallina I.Verd.A. dabenorisana Van Jaarsv.A. dewinteri GiessA. dinteri A.BergerA. distans Haw.A. erinacea D.S.HardyA. falcata BakerA. fouriei D.S.Hardy & GlenA. gerstneri ReynoldsA. gracilis Haw. var. decumbens ReynoldsA. haemanthifolia A.Berger & MarlothA. hardyi GlenA. inconspicua PlowesA. keithii ReynoldsA. khamiesensis PillansA. krapohliana MarlothA. longistyla BakerA. meyeri Van Jaarsv.A. microcantha Haw.A. modesta ReynoldsA. monotropa I.Vcrd.A. namibensis GiessA. pearsonii SchonlandA. peglerae SchonlandA. petrophila PillansA. picti$olia D.S.HardyA. pillansii L.GuthrieA. polyphylla Schonland ex PillansA. pratensis BakerA. prinslooi IVerd. & D.S.HardyA. pruinosa ReynoldsA. ramosissima PillansA. reitzii Reynolds var. reitziiA. reitzii Reynolds var. vernalis D.S.HardyA. reynoldsii Let tyA. saundersiae (Reynolds) ReynoldsA. simii Pole-EvansA. sladeniana Pole-EvansA. soutpansbergensis I.Verd.A. striata Haw. ssp. komaggasensis (Kritzinger & Van Jaarsv.)

Glen & D.S.HardyA. thompsoniae Groenew.A. thomcroftii Pole-EvansA. vandermerwei ReynoldsA. viridiflora ReynoldsA. vogtsii ReynoldsA. vossii ReynoldsA. vryheidensis Groenew.Astroloba herrei UitewaalGasteria batesiana G.D.RowleyG. baylissiana RauhG. bicolor Haw. var. liliputana (Poelln.) Van Jaarsv.G. croucheri (Hook.f.) BakerG. ellaphieae Van Jaarsv.G. glomerata Van Jaarsv.G. nitida (Salm-Dyck) Haw. var. armstrongii (Schonland) Van

Jaarsv.G. rawlinsonii Oberm.

RVERVEVERVRVVVVVIVEEEEVIEEVVEEEVEEEVVEVRVR

G. vlokii Van Jaarsv.Haworthia archeri W.F.Barker ex M.B.Bayer var. archeriH. archeri W.F.Barker ex M.B.Bayer var. dimorpha M.B.BaycrH. blackburniae W.F.BarkerH. bruynsii M.B.BayerH. comptoniana G.G.Sm.H. emelyae Poelln. var. emelyaeH. emelyae Poelln. var. mult[folia M.B.BaycrH. fasciata (Willd.) Haw.H. floribunda Poelln.H. graminifolia G.G.Sm.H. heidelbergensis G.G.Sm.H. kingiana Poelln.H. koelmaniorum Oberm. & D.S.HardyH. limifolia Marloth var. gigantea M.B.BayerH. limifolia Marloth var. limt$olia *H. limifolia Marloth var. ubontboensis (I.Verd.) G.G.Sm.H. lockwoodii ArchibaldH. magnifica Poelln. var. atrofrrsca G.G.Sm.H. magnifica Poelln. var. major (G.G.Sm.) M.B.BaycrH. magnifica Poelln. var. paradoxa (Poelln.) M.B.BaycrH. marginata (Lam.) StearnH. maughanii Poelln.H. mcmurtryi C.L.ScottH. mirabilis (Haw.) Haw. ssp. badia (Poclln.) M.B.BaycrH. mirabilis (Haw.) Haw. ssp. mundula (G.G.Sm.) M.B.BaycrH. mutica Haw.H. nortieri G.G.Sm. var. globosiflora (G.G.Sm.) M.B.BayerH. parksiana Poelln.H. pehlemanniae C.L.ScottH. poellnitziana UitewaalH. pubescens M.B.BayerH. pygmaea Poelln.H. retusa (L.) Duval var. dekanahii (G.G.Sm.) M.B.BaycrH. serrata M.B.BayerH. sordida Haw.H. springbokvlakensis C.L.ScottH. starkiana Poelln. var. lateganiae (Poclln.) M.B.BayciH. truncata SchonlandH. wittebergensis W.F.BarkerH. woolleyi Poelln.Poellnitzia rubrijZora (L.Bolus) Uitewaal

APOCYNACEAEI Adenium swazicum Stapf *V Pachypodium namaquanum (Wyley ex Harv.) Wclw.

ASCLEPIADACEAEIRVRVRRRRKKRRRRRVI

Ceropegia barbata R.A.DyerC. cancellata Rchb.C. cimiciodora Oberm.C. decidua E.A.Bruce ssp. pretoriensis R.A.DycrC. occidentalis R.A.DyerC. radicans Schltr. ssp. radicansC. verruculosa (R.A.Dyer) D.V.FicldCynanchum meyeri (Decne.) Schltr.Duvalia parviflora N.E.Br.D. pillansii N.E.Br. var. albanica N.E.Br.D. pillansii N.E.Br. var. pillansiiHoodia dregei N.E.Br.H. juttae DinterH. officinalis (N.E.Br.) Plowes ssp. delactiurta (Dintcr) BruynsH. pilifera (L.f.) Plowes ssp. annulata (N.E.Br.) BruynsH. pilifera (L.f.) Plowes ssp. pilifcraH. pilifera (L.f.) Plowes ssp. pillansii (N.E.Br.)H. ruschii Dinter

181

RIRRVVREIRRExExRRKKIRVIRVRVVKR

VKKRVRRRRR

RRRRRRRRRVVRIVKIRRRRRKR/VV/ERV

H. triehneri (Nel) BruynsHuernia echidnopsioides (L.C.Leach) L.C.LeachH. hallii E. & B.M.LambH. humilis (Masson) Haw.H. hystrix (Hook-f.) N.E.Br. var. pan&a L.C.LeachH. kennedyana LavranosH. longii PillansH. nouhuysii I.Verd.H. pendula E.A.BruceH. plowesii L.C.LeachH. praestans N.E.Br.H. thudichumii L.C.LeachH. witzenbergensis C.A.Liickh.Lavrania haagnerae PlowesNotechidnopsis columnaris (Nel) Lavranos & BleckOrbea irrorata (Masson) L.C.Leach0. lepida (Jacq.) Haw.0. macloughlinii (I.Verd.) L.C.Leach0. maculata (N.E.Br.) L.C.Leach0. paradoxa (IVerd.) L.C.Leach0. pulchella (Masson) L.C.Leach0. rangeana (Dinter & A.Berger) L.C. Leach0. speciosa L.C.Leach0. woodii (N.E.Br.) L.C.LeachOrbeanthus conjunctus (A.C.White & B.Sloane) L.C.Leach0. hardyi (R.A.Dyer) L.C.LeachOrbeopsis albocastanea (Marloth) L.C.Leach0. gerstneri (Letty) L.C.Leach ssp. elongata (R.A.Dyer)

L.C.Leach0. gerstneri (Letty) L.C.Leach ssp. gerstneri0. knohelii (E.Phillips) L.C.Leach0. tsumebensis (Oberm.) L.C.LeachPachycymbium lancasteri LavranosP. ubomboense (IVerd.) M.G.Gilbert *Pectinaria articulata (Aiton) Haw. ssp. articulataP. articulata (Aiton) Haw. ssp. borealis BruynsP. longipes (N.E.Br.) BruynsPiaranthus barrydalensis MeveQuaqua armata (N.E.Br.) Bruyns ssp. arenicola (N.E.Br.)

BruynsQ. armata (N.E.Br.) Bruyns ssp. maritima BruynsQ. j?amesii (Pillans) BruynsQ. inversa (N.E.Br.) Bruyns var. cincta (C.A.Luckh.) BruynsQ. inversa (N.E.Br.) Bruyns var. inversaQ. linearis (N.E.Br.) BruynsQ. multiflora (R.A.Dyer) BruynsQ. parviflora (Masson) Bruyns ssp. bayeriana BruynsQ. pruinosa (Masson) BruynsStapelia baylissii L.C.LeachS. clavicorona I.Verd.S. divaricata MassonS. erectiflora N.E.Br. var. prostratiflora L.C.LeachS. glabricaulis N.E.Br.S. immelmaniae PillansS. macowanii N.E.Br. var. macowaniiS. obducta L.C.LeachS. paniculata Willd.S. pearsonii N.E.Br.S. peglerae N.E.Br.S. praetermissa L.C.Leach var. luteola L.C.LeachS. praetermissa L.C.Leach var. praetermissa L.C.LeachS. remota R.A.DyerS. rubiginosa NelS. scitula L.C.LeachS. tsomoensis N.E.Br.Stapeliopsis breviloba (R.A.Dyer) Bruyns

R S. exasperata (Bruyns) BruynsE S. neronis PillansR S. pillansii (N.E.Br.) BruynsV S. saxatilis (N.E.Br.) Bruyns ssp. stayneri (M.B.Bayer) BruynsR S. urniflora LavranosWV Tridentea pachyrrhiza (Dinter) L.C.LeachR 7’. virescens (N.E.Br.) L.C.LeachWV Tromotriche herrei (Nel) L.C.LeachR T. longii (C.A.Luckh.) L.C.LeachR T. ruschiana (Dinter) L.C.LeachR T. thudichumii (Pillans) L.C.Leach

ASPHODELACEAER Bulbine brunsvigiifolia BakerR B. diphylla Schltr. ex Poelln.R B. fallax Poelln.R B. francescae G.Will. & Baijnath ined.V B. haworthioides B.Nord.R B. louwii L.I.HallE B. margarethae L.I.HallK B. monophylla Poelln.R B. orchioides Drege ex Poelln.R B. striata Baijnath & Van Jaarsv.V B. wiesei L.I.Hall

ASTERACEAEK Othonna abrotanifolia (Harv.) DruceR 0. armiana Van Jaarsv.K 0. cacalioides L.f.R 0. hallii B.Nord.K 0. lepidocaulis Schltr.WV 0. rechingeri B.Nord.K Senecio scaposus DC. var. addoensis (Compton) G.D.RowlcyK S. haworthii (Sweet) Sch.Bip.K S. medley-woodii Hutch.K S. muirii L.Bolus

BRASSICACEAER Heliophila eximia Marais

CRASSULACEAEKKKIIKRRKRKRRR

RExKRRRKR/VRR

Adromischus bicolor HutchisonA. cooperi (Baker) A.BcrgerA. cristatus (Haw.) Lem. var. zeyheri (Harv.) TA. diabolicus ToelkenA. fallax ToelkenA. humilis (Marloth) Poelln.A. liebenbergii HutchisonA. mammillaris (L.f.) Lem.A. marianiae (Marloth) A.Berger var. murianiaeA. nanus (N.E.Br.) Poelln.A. phillipsiae (Marloth) Poelln.A. schuldtianus (Poelln.) Poelln. ssp. juttae (Poclln.) ToelkenA. subviridis ToelkenCotyledon orbiculata L. var.j7amgmrzii (Schiinland & Baker f.)

ToelkenC. tomentosa Harv. ssp. ladismithiensis (Poelln.) ToelkcnCrassula alcicornis SchonlandC. alpestris Thunb. ssp. massonii (Britten & Baker f.) ToelkcnC. arborescens (Mill.) Willd. ssp. undulatijolia ToelkcnC. aurusbergensis G.Will.C. ausensis Hutchison ssp. giessii (Friedrich) ToelkenC. brachystachya ToelkenC. brevifolia Harv. ssp. psammophilu Toe1C. decidua SchonlandC. elsieae Toelken

ken

182

R

VWVRKR

RRRRRExIR

RKRRRRRRRVRRKERRRR

C. exilis Harv. ssp. exilisC. garibiraa Marloth & Schonland ssp. glabra ToelkenC. latibracteata ToelkenC. luederitzii SchonlandC. multiceps Harv.C. multiflora Schonland & Baker f. ssp. leucantha (Schonland &

Baker f.) ToelkenC. namaquensis Schonland & Baker f. ssp. comptonii (Hutch. &

Pillans) ToelkenC. planifolia SchonlandC. plegmatoides FriedrichC. qoatlhambensis Hargr.C. rupestris Thunb. ssp. commutata (Friedrich) ToelkenC. rupestris Thunb. ssp. mamierana (H.E.Huber & Jacobsen)

Toe1 kenC. sericea Schonland var. velutina (Friedr.) ToelkenC. sladenii SchonlandC. socialis SchonlandC. streyi ToelkenC. subacaulis Schonland & Baker f. ssp. subacaulisC. subulata L. var. hispida ToelkenC. susannae Rauh & FriedrichC. thunbergiana Schult. ssp. minutiflora (Schonland & Baker f.)

ToelkcnC. tuberella Toe1 kenCrassula vaginata Eckl. & Zeyh. ssp. minuta ToelkenC. vestita Thunb.Tylecodon albiflorus BruynsT atropurpureus BruynsT. bayeri Van Jaarsv.T. boddleyae Van Jaarsv.T. decipiens ToelkenT. ellaphieae Van Jaarsv.T -fragilis (R.A.Dyer) ToelkenT. hirtifolius (W.F.Barker) ToelkenT. kritzingeri Van Jaarsv.T. occultans (Toelken) ToelkenT. singularis (R.A.Dyer) ToelkenT. suffultus Bruyns ex ToelkenT. sulphureus (Toclken) ToelkenT. tenuis (Toclken) BruynsT. torulosus Toelken

EUPHORBIACEAER/VVKRVVRRExRRVVKExVR/VVIRRE

Euphorbia albertensis N.E.Br.E. albipollin~fera L.C.LeachE. artijolia N.E.Br.E. baliola N.E.Br.E. barnardii A.C.White, R.A.Dyer & B.SloaneE. bayeri L.C.LeachE. berotica N.E.Br.E. brakdamensis N.E.Br.E. brevirama N.E.Br.E. bruynsii L.C.LeachE. cibdela N.E.Br.E. clavigera N.E.Br.E. clivicola R.A.DyerE. corymbosa N.E.Br.E. crassipes MarlothE. cylindrica A.C.White, R.A.Dyer & B.SloaneE. decepta N.E.Br.E. fasciculata Thunb.E. friedrichiae DinterE. globosa (Haw.) SimsE. grandialata R.A.DyerE. groenewaldii R.A.Dyer

RV/ERKKERKRRRKRRVV/EKRREKRWEIKEKV/EKRRRRKVRVVRRVKRVRR

E. hallii R.A.DyerE. hopetownensis NelE. hottentota MarlothE. inomata N.E.Br.E. insarmentosa P.G.Mey.E. jansenvillensis NclE. kaokoensis (A.C.White, R.A.Dyer & B.Sloane) L.C.LeachE. keithii R.A.DyerE. knobelii LettyE. lavrani L.C.LeachE. ledienii A.Berger var. dregei N.E.Br.E. louwii L.C.LcachE. lumbricalis L.C.LeachE. malevola L.C.Leach ssp. bechuanica L.C.LcachE. marlothiana N.E.Br.E. meloformis AitonE. monteiroi Hook.f. ssp. hizrrllll~el~e}l.~i.s B.Nord.E. namusk1uftensi.s L.C.LeachE. nesemannii R.A.DyerE. obesa Hook.f.E. ornithopus J acy.E. otjipembana L.C.LeachE. oxystegia Boiss.E. pedemontana L.C.LeachE. pentops A.C.White, R.A.Dyer & B.SloancE. perangusta R.A.DyerE. planiceps A.C.White, R.A.Dyer & B.SloancE. polycephala MarlothE. pseudoglobosa MarlothE. quadrata NelE. restituta N.E.Br.E. restricta R.A.DyerE. rowlandii R.A.Dycr “’E. rudolfii N.E.Br.E. schoenlandii PaxE. sekukuniensis R.A.DyerE. symmetrica A.C.Whitc, R.A.Dyer C! B.SloancE. tortirama R.A.DyerE. umfoloziensis PcckoverE. vaalputsiana L.C.LeachE. valida N.E.Br.E. venenata MarlothE. versicolores G.Will.E. waterbergensis R.A.DyerE. woodii N.E.Br.E. zoutpansbergensis R.A.Dycr

GERANIACEAEI Pelargonium crassipcs HaW.R P. desertonlm VorsterR P. hystrix Harv.R P. tort&sum E.M.MaraisK Sarcocaulon herrei L.BolusK S. inerme RehmR S. peniculinum Moffctt

PASSIFLORACEAER Adenia fruticosa Burtt Davy ssp. sirrtplict’folirr W.J.dc WiltEx A. natalensis W.J.de WildeR A. pechuelii (En@.) Harms

PORTULACACEAER Anacampseros bayeriana S.A.HammerR A. comptonii PillansR A. dinteri SchinzR/V A. filamentosa (Haw.) Sims ssp. filanterttosa

183

R A. filamentosa (Haw.) Sims ssp. tomentosa (A.Berger)

RRRRRIRRRRKR

GerbauletA. herreana Poelln.A. karasmontana DinterA. lanceolata (Haw.) Sweet ssp. lanceolataA. lanceolata (Haw.) Sweet ssp. nebrownii (Poelln.) GerbauletA. papyracea E.Mey. ex Fenzl ssp. papyraceaA. yuinaria E.Mey. cx FenzlA. recurvata Schonland ssp. buderiana (Poelln.) GerbauletA. recurvata Schonland ssp. minuta GerbauletA. recurvata Schonland ssp. recurvataA. rhodesica N.E.Br. *A. rufescens (Haw.) SweetA. ruschii Dinter ex Poelln.

RRRV

A. scopata G.Will.A. subnuda Poelln. ssp. lubbersii (Bleck) GerbauletAvonia mallei G.Will.Portulaca trianthemoides Bremek.

VITACEAE

R Cyphostemma bainesii (Hook.f.) Desc.K C. hardyi RetiefR C. juttae (Dinter & Gilg) Desc.R C. uter (Exe11 & Mendonca) Desc. ”

ZYGOPHYLLACEAE

R Zygophyllum tereti$olium Schltr.

184

Annex 9

Threatened succulents of ZimbabweCompiled by Craig Hilton-Taylor.

The data presented here were extracted from Kimberley (1991) and the World Conservation Monitoring Centrc (WCMC l993), with corrections basedon information from the ‘SARARES’ database and unpublished notes of M.J. Kimberley. All species are endemic unless noted by an asterisk (“).

status’Local Global

Taxon

I

RVRVVVRVREVIVIERVV/EV/E

VIER

ntV

ntV

VntIntnt

ntnt

AIZOACEAEDelosperma steytlerae L.BolusALOACEAEAloe ballii ReynoldsA. cameronii Hemsl. var. bandana ReynoldsA. chabaudii Schiinland var. verekeri Christian *A. hazeliana Reynolds *A. howmanii ReynoldsA. inyangensis ChristianA. lutescens Groenew. *A. munchii Christian *A. musapana ReynoldsA. ortholopha Christian & Milne-Redh.A. plowesii Reynolds *A. pretoriensis Pole-Evans *A. rhodesiana Rendle *A. sessiliflora Pole-Evans *A. suffidta Reynolds *A. tauri L.C.LeachA. wildii (Reynolds) Reynolds

APOCYNACEAEAdenium multiflorum Klotzsch *Pachypodium saundersii N.E.Br. *

ASCLEPIADACEAEE nt Hoodia currorii (Hook.) Decne. ssp. lugardii (N.E.Br.)

Bruyns *R Huernia longituba N.E.Br. ssp. cashelensis L.C.Leach

& PlowesR H. occulta L.C.Leach & PlowesR Orhea umbracula (M.D.Hend.) L.C.LeachE nt Tavaresia harklyi (Dyer) N.E.Br. *

CRASSULACEAER Kalanchoe velutina Welw. ex Britten ssp.

chimanimanensis (R.Fern.) R.Fern.R K. lohata R.Fern.R K. wildii Raym.-Hamet ex R.Fern.

EUPHORBIACEAER nt Euphorbia confinalis R.A.Dycr ssp. conjintr1i.s ‘.V E. confinalis R.A.Dyer ssp. rhodcsica L.C.LeachV nt E. cooperi N.E.Br. ex A.Bergcr var. calidicolrr

L.C.Leach *RVIERERREEVRRV/EVE

ntnt

Intntnt?

R

nt

nt

E. davyi N.E.Br. ”E. decidua P.R.O.Bally & L.C.Lcach ‘I’E. dissitispina L.C.LeachE. fortissima L.C.Lcach ”E. gossypina Pax var. gosg@lrr “’E. guerichiana Pax ”E. halipedicola L.C.Leach ‘I’E. li~idiflorn L.C. Leach “’E. memoralis R.A.DyerE. rowlandii R.A.Dyer ‘!’E. rugosijlora L.C.LeachE. trichadenia Pax ”E. wildii L.C.LeachMonadenium lugardae N.E.Br. “’

PASSIFLORACEAER R Adenia jfuticosa Burtt Davy ssp. .simplic~‘i~lia W.J.dc

Wilde :g

VV nt

A. karihac~~sis W.J.de WildeA. spinosa Burtt Davy ‘!’

’ Column one indicates the conservation status of each taxon withinZimbabwe; the global status is indicated in column two, cxccpt forendemics where the global status is the same as that for the country. Theglobal status was obtained from the WCMC (1993) and updated based oninformation from the ‘SARARES’ database.

’ The taxon name given generally follows the nomenclature used inArnold and De Wet (1993). however, there are differences whet-o WC

have followed taxonomic treatments published subsequent to thosereferred to in Arnold and DC Wet (1993) or taxonomic tre:itments notfollowed by the National Herbarium in Pretoria, South Africa (PKE).The abbreviations of author names are according to Brummitt and Powell(1992). The taxa are arranged in alphabetical order by family then !+~us.The family names used are those most commonly used. Families whichare often given alternative names are (name used here/alternative):Aloaceae/Asphodelaceae, Aizoaceae/Mesembry~~nIhem~rceac, andAsteraceae/Compositae.

185

Annex10

Threatened succulents of India

Information provided by C. R. Babu and Meena Singh, reviewed by S. Karthikeyan.

India has an estimated 15,000 vascular plant species including pteridophytes and about 5000 endemic flowering plants.The geographical position of India and its varied climatic conditions are factors which contribute to the diversity of itsflora. Many species are of economic or medicinal value and local societies traditionally have managed plant resourcessustainably; however dramatic population growth in the current century has placed an enormous strain on these naturalresources.

Destruction of forests for local fuel use, for timber by outside agencies, over-grazing by cattle and goats, quarryingand mining, development activity and reclamation of “unproductive” land have all lead to a decline in the country’sfloristic diversity. Forests have undergone varying degrees of transformation, ranging from degenerative changesbrought about by accelerated rates of soil erosion to complete replacement by monoculture. The effects of over-exploitation are manifested in increasing desertification of our lands and changing global climate patterns. Muchconservation work is called for in India, but regeneration of endemic flora in regions characterised by economic under-development and rural poverty will only succeed as long as it yields diverse and tangible benefits to inhabitants of theregion.

Here we present a conservation assessment and status report for India’s most threatened cacti and succulents. Basicsurvey and population monitoring, as well as conservation status reports, remain to be carried out for many taxa.

ASCLEPIADACEAEThe economic importance of the family includes the use of some species of Carahma such as C. c&4Zis, as greenvegetables and the tubers of Ceropegia species as food.

CeropegiaThe Indian subcontinent is one of the primary centres of genetic diversity for Ceropegia; here are about 45 indigenousspecies including both succulents with tuberous rootstocks and non-succulent herbaceous plants. The genus ranges fromhumid tropical regions to the temperate zone of the Himalayas with the chief centre of distribution in the Western Ghatmountains, from Maharashtra to Kerala. Most of the species are endemic to India and 27 species are endemic topeninsular India. Deforestation and overexploitation are among the many threats to these taxa causing some species,such as C. Zongifolia, C. wallichii, and C. macrantha in the sub-Himalayan tracts and Arvallis, to be on the verge ofextinction. C. bulbosa is almost extinct from Delhi and the adjoining areas, and species reported from sub-Himalayantracts have not been re-located in spite of repeated field trips to known localities. Immediate inventory, monitoring, andin situ conservation measures for these taxa are required along with ex situ conservation in botanic gardens (Babu 1994).

R

E

nt

nt

E

E

R

E

nt

R

nt

E

R

E

R

R

R

E

E

Ceropegia a ttenua taC. bamesiiC. ciliataC. ensifoliaC. evansiiC. fantasticaC. jkn briqera

C. huberiC. in termediaC. jainiiC. junceaC. lawiiC. maccaniiC. mahabaleiC. mediaC. noorjahanaeC. oculata

C. odorataC. omissa

Endemic to Maharashtra and Karnataka. Tubers are harvested for food.

Endemic to Tamil Nadu and Karnataka. Endangered due to habitat loss.

Endemic to Kerala and Tamil Nadu.


Endemic to Maharashtra. Tubers are edible. Endangered due to harvests from the wild.

Endemic to Goa and Karnataka.

Endemic to Karnataka and Tamil Nadu.

Endemic to Maharashtra. Endangered due to landslides.


Endemic to Maharashtra.

Endemic to Andhra Pradesh, Tamil Nadu, and Karnataka. The extract of the plant is used as a tranquilizer.

Endemic to Maharashtra. Endangered due to habitat destruction.





Endemic to Maharashtra and Kerala.

Endemic to Gujarat and Maharashtra. It is protected in the Amravati Tiger Reserve.

Endemic to Kcrala and Tamil Nadu.

186

E C. panchganiensisR C. pusillaR C. rollaeE C. sa hayadricaR C. santapaui

R C. spira lisE C. vincaefolia

Endemic to Maharashtra. It appears to be on the verge of extinction due to harvest of edible tubers from the wild.

Restricted to Karnataka, Kerala, and Tamil Nadu.


Endemic to Maharashtra. Endangered due to overcollection of its edible tubers.


Restricted to Andhra Pradesh, Karnataka, Kerala, and Tamil Nadu.

Endemic to Maharashtra. Endangered due to overcollection of its edible tubers.

CarallumaIn India there are 13 species of Caralluma most of which are confined to Peninsular India; 6 are endemic to PeninsularIndia and are found in most of the states located in the phytogeographical zone (Ahmedullah 1993). Populations aredeclining in size and distribution range due to overexploitation, mining, and deforestation. The habitats of this genus,particularly in the Western and Eastern Ghats and Vindhyan ranges, are fast disappearing due to mining activities.Inventory of the genus is urgently needed. Habitat protection will hopefully ensure the conservation of this unique groupof plants.

Rnt/RIRntRntKR

Caralluma edulisC. indicaC. lasianthaC. nilagirianaC. paucijloraC. procumbensC. stalagmiferaC. trunca to-corona taC. tu bercula taC. urn bella ta

Restricted to a small area in Rajasthan.Not threatened according to Babu (1994); Rare according to Singh (1994a); edible and emoycd by shepherd boys.

Described from a single clone and has not been found again in the wild.

Found only on a few remote hills of Tamil Nadu.Not threatened according to Babu (1994); threatened by proposed housing development according to Singh ( 1994a).Endemic to Karnataka.An edible species found on the lower tracts of the northern Himalayas grazed by sheep and goats.Threatened by habitat clearance.

BrachystelmaPeninsular India is the second centre of origin of the genus Brachystelma. Species are distributed in the hill ranges ofMadhya Pradesh, Orissa, Andhra Pradesh, Tamil Nadu, Kerala, Goa, Maharashtra, Gujarat, and Karnataka. Of the tenspecies found in India, nine are endemic to Peninsular India; the only species occurring outside this region is B.pawi$bms in the north-west (Ahmedullah 1993). The main threat to the species is quarrying and mining (Babu 1994).

Brachystelma bourneaeB. brevitubula turnB. cilia tumB. glabrumB. kolarensisB. maculatumB. rangachariiB. volubile

Endemic to Tamil Nadu.Endemic to Tamil Nadu.Endemic to Karnataka.Endemic to Andhra Pradesh.Endemic to Karnataka.Endemic to Karnataka and Tamil Nadu.Endemic to Coimbatore.Endemic to Andhra Pradesh.

DischidiaThree species are found in the Andaman Islands. Dischidia bengalcnsis and D. nimuzaria can be found extensively whileD. raffelsiana var. major is Rare.

--HoyaThe genus Hoya is widely distributed in the temperate regions of the Himalayas where species are under threat due todeforestation. Since forty percent of the forest cover in the Himalayas has been lost, substantial numbers of populationsof Himalayan Hoya species have also been lost. The remainder of these species requires immediate conservationmeasures. H. wightii is an endemic species of Peninsular India but is likely to become extinct when its habitat issubmerged upon completion of the Idukki Hydroelectric dam.

CRASSULACEAEThe Himalayas constitute one of the centres of diversity for the genus Sedum. The genus requires inventory andmonitoring and critical taxonomic assessment before its conservation status is assessed. KaZarzchoe clzeI.likorlCIerl~si,s, anendemic of Eastern Ghats (Visakhapatnam Hills), is also endangered due to habitat destruction.

187

CUCURBITACEAEA few Indian species with tuberous roots can be considered semi-succulent. Of these the genera C~~~lloccrrpus, Kedrostis,and Momordica are widely distributed in Peninsular India. Coralocarpus epigaeus tubers are of medicinal value and thespecies is threatened by over-exploitation. The fruits of Momordica dioica are edible and attempts are being made tocommercialise this species.

EUPHORBIACEAEWithin the genus Euphorbia, 19 Indian species show succulence. Of these 16 are dendroid and three are geophytic. AllIndian Jatropha species are semi-succulent.

The cactoid euphorbias are the major elements of dry slopes and cliffs in all the hill ranges and the Himalayas up to4000 m. In the dry hill tracts of Peninsular India they comprise the climax formation. These cactoid euphorbias includetree euphorbias with fleshy stems with or without leaves as well as pencil euphorbias. The taxonomy of these species isnot yet fully worked out and needs further attention. Inventory and reclassification is also needed for the three geophyticEuphorbia spp.

Some of the tree euphorbias have been utilised as renewable sources of energy and also in making toys. This has ledto depletion of populations in the Himalava and other hill ecosvstems.

J r’

Euphorbia antiquorum

E. atotoE. harnhartii

E. ca ttimandooE. cauduc(foliaE. corrigioloidesE. epiphylloides

E. fitsiformis

E. khandallensis

E. mayurnathaniiE. neriijblia

E. nivuliaE. panchagmiensis

E. royleana

E. santapauii

E. senguptaeE. swan-holmcsiae

E. tortilis

E. vajravelui

This species mostly grows in mixed populations with E. tortilis (see below). It is threatened by collection for fuel andland reclamation.

A semi-succulent species of coastal regions; apparently rare.Very rare. This species has disappeared in the wild on the Indian mainland. Small populations still exist in the

Andaman Islands but are under threat as habitat is cleared for cultivation by settlers.Very rare. Found only in a small area in Andhra Pradesh; habitats have been cleared for cultivation.Common in dry rocky areas of Rajasthan, Gujarat, and Andhra Pradesh.Endemic to Peninsular India. It has a tuberous rootstock and is herbaceous.Very rare. Although the habitat of this species is in a reserved forest the population is declining for unknown reasons.

Serious conservation attention is required for this species.Restricted to north-western sub-Himalayan tracts; it is endangered due to overexploitation of its medicinally

important tuberous rootstock (Babu, in litt, 1994). Four or five sites have been located within a 300 km radius(Singh, in litt. 1994b).

Endemic to Maharashtra where it is restricted to a small area in Khandala and Lonavala. A hcrbaceous species withtuberous rootstock; its habitat is very stony and poor for cultivation, but land is being reclaimed for farming andholiday retreats.

Described from a goup of three isolated trees and now extinct.This species does i lot seem to be threatened as populations in restricted areas are doing well. This spccics is

commonly seen as a hedge plant in Maharashtra.Few wild populations of this species remain.Endemic to a small area in the Panchgani Hills, Maharashtra. A herbaceous species with cylindric underground

rootstock. Threatened by the conversion of its habitat into a large fairground.A species of the sub-Himalayan tracts. Habitat has shrunk due to land clearing but is still common in various

locations.Endemic to Tamil Nadu. A shrub or tree with fleshy leaves. This species is endemic to Mount Agasthaya on the

border between Kerala and Tamil Nadu, recently declared a Biosphere Reserve.Endemic to Kurnool. An erect herb.Very rare; a recently described species which is threatened by periodic river flooding washing away the seeds so that

the species cannot regenerate. Relocation of plants needs to be considered.This species is commonly seen as a hedge plant, but has been reduced in its natural habitats by felling for fuel and by

land reclamation.Rare; a recently described species, found on the lower slopes of the Western Ghats. Threatened as habitats arc hcing

cleared to make way for tea plantations.

188

Annex 11

Threatened succulents of Mexico

Categories of threat are given by SEMARNAP: P = in danger of extinction (en Peligro de Extincibn), A = threatened (amenazada), R = rare @-a).The corresponding IUCN categories are as follows: P = Endangered, A = Vulnerable, R = Rare. An asterisk (*) indicates an endemic taxa.

AGAVACEAEAgave bracteosaA. chiapensisA. congestsA. dasylirioidesA. guiengolaA. gypsophilaA. impressaA. kewensisA. luridaA. nizandensisA. ornithobromaA. parrasanaA. parvifloraA. peacockiiA. polianthiJoraA. titanotaA. Victoria-reginaeA. vizcainoensisBeaucarnea gracilisB. goldmaniiB. hiriartiaeB. plia bilisB. purpusiiB. recurva taB. strictaBeschorneria albifloraB. calcicolaB. tu bifloraB. wrigh tiiDasylirion palaciosiiD. longissimumFurcraea bendinghausiiF. macdougalliiManfreda brunneaM. guerrerensisM. longifloraM. nanchititlensisM. planifoliaM. potosinaPolianthes densifloraP. howardiiP. longifloraP. palustrisP. platyphyllaYucca endlichianaY grandifloraY lacandonicaY queretaroensis

ASCLEPIADACEAEAsclepias macvaughii

CACTACEAEAporocactus leptophisA. flagelliformisAriocarpus agavoidesA. bravoanus

AR*RA*A”RA*R*P”P*RR*AR*AR*P”R”A*A*A*A”A*AA*RR”RR”R*AA*P*AR*AA*R*R”R*R*R*R*R*RR’”AR

R

P”R”P”R””

A. fissura tusA. kotschoubeyanusA. scapharostrusA. trigonusAstrophytum asteriasA. capricorneA. ornatumA. myrios tigmaA. hintoniiA. ritteriBackebergia militarisCephalocereus nizandensisC. senilisCoryphantha delica taC. durangensisC. elephan tidensC. glanduligeraC. gra cilisC. grataC. green woodiiC. odorataC. poselgerianaC. pseudoechinusC. pulleinianaC. ramillosaC. retusa var. melleospinaC. sch warzianaC. sulcata var. nickelsiaeC. werdermanniiCryptocereus anthonyanusEchinocactus grusoniiE. parryiE. platyacanthusEchinocereus adustusE. bristoliiE. delaetiiE. freuden bergeriiE. knippelianusE. MauiE. leucanthusE. lindsayiE. longisetusE. nivosusE. palmeriE. poselgeriE. pulchellusE. reichenbachii var. fitchiiE. schmolliiE. sciurusE. stolonifelusE. subinermisE. wein bergiiEchinomastus erectocentra

var. acunensisE. in tertextusE. mariposensi

189

A*A”P”A”P*A”A:‘:A*R*A*R*R””A”R*R*AA””P*R”R”R*A”R*A*AR”R”AP*AP”A”

Pr*A””RA”A:‘:A”A”R”P”R””R”PPA*A*P”R”R*R”R”

PA”A”

E. unguispinus var. dllrangerlsisE. unguispinlls var. lalliE. unguispinus var. unguispimlsE. warnockiiEpiphyllum c11~~‘ioc’ar~~illrlrEpithelantha bokciE. micromerisEscobaria aguirTc>anrrE. asperispinaE. chafleyiE. laredoiE. roseanaFerocactus clliysnc~crrltlrr(,sF. cylindraccrrsF. /zaer~lntacarltllrr.sF. johnstonianusF. pilosrrsF. reppenhagenniF. totv~lsel1ciia~zrr.s var. ton~ilsc~~~ii~r~1rr.sF. viridescensGeohintonia mexicanaHamatocactus C’r~~s.sillCrf?rrrtrl~sH. uncinatusLeuchtenbergia princi$isLophocereus schottii

var. mieckleyanusLophophora di#ll.saL. williamsiiMammillaria a1bica~i.sM. albicomaM. angelensisM. annianaM. aureicepsM. aureilanataM. aurihanlataM. backebcrgianaM. barrmiiM. bciscliiM. bloss&4~ia~l aM. bocasanaM. bon?bJxirlaM. booliiM. candidaM. capensisM. carmenaeM. caiT*etiiM. cclTalboaM. coahrrilensisM. crucigcraM. dehcrdtiana var. delrcw’tiantrM. dcherdtiana var. dodsorliiM. cjrjia?ltllOC’eIlt1.(~11M. dl1qfc)rmisM. crectacanthaM. ~~~~er~~~a~~~~i~~~~~~M. fittkaui

AA”.R”RAZkARR:‘:A*A:”R*R*

A*RR’-KSA*R*AA$:R*A*A>‘:A:‘:

R”:A:‘:PtR>‘-AZkR*:R*A:‘:Ad:R:”R:i:R”’R”:R”A:‘:R*R:”A+R”PZ’:R3<R’j:A -‘.R “.R *A:;:R:”R3:A”.R -‘:R”.

M. gaumeriM. glareosaM. goodridgiiM. grusoniiM. guelzowianaM. guerreronisM. hahnianaM. haleiM. heidiaeM. hernandeziiM. herreraeM. hertrichianaM. huitzilopochtliM. humboldtiiM. insularisM. johnstoniiM. klissingianaM. knippelianaM. kraehenbuehliiM. laui var. laui- var. dasyacantha- var. discataM. lentaM. lindsayiM. longifloraM. longimammaM. magnificaM. maritimaM. marksianaM. mathildaeM. matudueM. melaleucaM. mercadensisM. meyruniiM. microheliaM. miegiunaM. moellerianaM. multidigita tuM. nanaM. napinaM. neopalmeriM. oteroiM. painteriM. parkinsoniiM. pectin~feraM. peninsularisM. pennispinosaM. perezdelarosaeM. pilcuyensisM. pilispinuM. plumosaM. pondiiM. pringleiM. pubispinaM. repenhageniiM. rettigiana

R:‘:R:k

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M. roseoalbaM. rubrograndisM. saboaeM. san-angelensisM. sanchezmejoradaeM. schiedeanaM. sch warziiM. senilisM. setispinaM. sleviniiM. solisioidesM. stella-de-tacubayaM. surculosaM. tayloriorumM. tepexicensisM. theresaeM. tonalensisM. varieaculeataM. weingartianaM. wiesingeriM. xaltianguensisM. yaquensisM. vucu tensisM. zeilmannianaM. zephymn thoidesMelocactus dawsoniiM. delessertianusM. ruestiiMitrocereus filvicepsMorangaya pensilisNopalxochia macdougalliiN. phyllanthoidesObregonia denegriiOpuntia an teojoensis0. arenaria0. bravoana0. excelsa0. rosarica0. santamariaOrtegocactus ma‘cdougalliiPachycereus guumeriPelecyphora uselliformisP. strobiliformisPeniocereus cuixmalensisP. fosterianusP. greggiiP. lazaro-cardenasiiP. maculatusP. marianusP.P.

tepalcatepecanuszopilotensis

Pilosocereus cometesPterocereus gaumeriSelenicereus anthonyanusS. a tropilosusStenocactus coptonogonus

R”R”

A*

P”

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S. sulphurz~rrsStenocereus C’12uC’Nl(I~~C’Il.si.sS. erucaS. martineziiStrombocactus discijormisThelocactus bicolor WI-. bolansisT. hasti/&T. ~~etcroc‘l?l.c~~zlrrsT. leucanthrrs var. ehrenbqiiT. macdoweliiT. rincon~~nsis WT. nidulattsT. sch warziiT. tulcnsisTurbinicarpus gautiiT. gielsdor-fianusT. h oferiT. lauiT. lop~lc~pllol.~~i~l~~.sT. mandragoraT. pseudonlnci.~~c.lii’lL’T. pscor(d~~~~ectirlatIrsT. saueriT. scll~~liedi~‘kCN~ll(.sT. s~1lr?liec~i~kc)allrls VU-. gracilisT. subter-rzrnerrsT. swobodacT. valdezianusT. vicreckiiT. ysa belae

CRASSULACEAEEcheveria amphoralisE. elegansE. lauiE. longissinta var. aztatlcnsisE. longissima VU-. longissimaE. moraniiE. prllp1’s”‘-ll’7l

E. sctosa VX. c’iliataE. setosa WI-. deminuataE. setosa VW. mirrorE. setosa var. otcrwE. setosa var. sctosGraptopetalum grrrnlk~G. macdougalliiSedum fi*utcscensS. pla~phyllrims. sll~weolms

s . tol*lllosllr?l

FOUQUIERIACEAEFouquieria fasciculataF. leonilacjF. ochotcrenacF. purpusiiF. shre\ei

R $R:;:

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190

Annex 12

Mexican succulent nurseriesI.T.E.S.M.-Campus QueretaroHenry Ford 10, Parques Industriales,Queretaro 76130 Qro.Tel: 91 (42) 17 38 28Fax: 91 (42) 17 38 23

Quinta Las CamelinasCarr. Cordoba-Fortin de las Flores, Km 133Fortin de las Flores, Ver.Tel: 91 (271) 308 08

Ranch0 La Joya, S.P.R. De R.I.Domicilio conocido,Col. Juan Ubera,Atlixco, Pue.Tel: 91 (244) 532 99Fax: 91 (244) 534 91

Orquideario De BajaCarr. Transpeninsular, Km 16 y Calle No. 5Ejido “El Centenario”,Baja California Sur, B.C.S.

Quinta Fernando Schmoll Vivero DeQuinta s/n.Cadereyta de Montes, Qro.

Vivero De Cactus Y Suculentas De CoahuilaLote no 5, Ex-Hacienda Los CarritosKm 5 de la Carretera a MexicoSaltillo, Coah.

Can Te, A.C.Mesones 71Col. Centro,San Miguel de Allende37700 Gto.Tel: 91 (465) 229 90Fax: 91 (465) 240 15

Follaje Y OrnatoBarrio de San Martinito s/nTlaxcalancingo Cholula, Pue.Tel: 91(22) 49 72 64

La Flor De CatemacoS.P.R. de R.L.de C.V.Carr. Catemaco-Coyame, Km 9.570607 Catemaco, Ver.Tel: 91 (294) 303 25Fax: 91 (294) 305 68

Orquideas Jardin Rio VerdeCalle de Arista 20151200, Valle de Bravo, Edo. de Mexico,Apartado Postal 69Tel: 91 (726) 202 89Fax: 91 (726) 209 11

Plantas y Excursiones de California SurCalle 12 y Ave. 10 s/nCol. LindavistaFortin de la Flores, Ver.

Mic-CactiPlantas, Cactaceas y SuculentasAve. San Jose, Lte. 8 Mza VllCo1 Granjas Residenciales de TequisquiapanTequisquiapan, Qro.

Monte ObscureEjido El PalmarCongregation Monte ObscureMpo. Emiliano Zapata, Ver.Tel: 91 (271) 464 99

Centro De Reproduction De Cactaceas “El Copal”Ejido Banco National AgrarioMapimi, Dgo.

Jose Raul Gomez Perez Vivero De Arboles, S.A. De C.V.

Miguel Rebolledo 4 Lope de Vega 791

Coatepec Jardines de1 Bosque

91500 Ver. 44520 Guadalajara, Jal.

191

AguascalientesJardin Botanic0 de1 Parque “HeroesMexicanos”Domicilio ConocidoAguascalientes, Ags

CampecheJardin Natural Forestal C.E.F“Ing. Eduardo Sangri Serrano”Carr. Fed. NO. 186 VillahermosaEscarcega Km. 292Escarcega, Campeche 24350

Jardin Botanic0 “Xmuch-Haltun”Calle 8-49Campeche, Campeche

CoahuilaJardin Botanic0 “Gustav0 AguirreBenavides”Universidad Autonoma Agraria“Antonio Narro”

Apdo. Postal 554Saltillo, Coahuila 25000

Jardin Botanic0 “Jerzy Rzedowski”Univ. Aut. Agraria “Antonio Narro”Unidad LagunaCarr. A Santa Fe y PerifericoApartado Postal 94 Carro 2Torreon, Coahuila

C hiapasJardin Botanic0 “Dr. Faustino Miranda”Inst. de Historia Natural de ChiapasDepartamento de Botnica5a. Norte Oriente Apdo. Postal 6Tuxtla-Gutierrez, Chiapas 29000

Orquidiariodel Consejo Protector de LaNaturaleza5a. Norte OrienteA. P. 391 Tuxtla-GutierrezChiapas 29000

Jardin Botanico “San Cristobal”Centro de Investigaciones Ecologicas de1SuresteCarretera Panamericana y Periferico SurSan Cristobal de Las Casas, Chiapas 29290

Annex 13

Mexican botanic gardens

ChihuahuaJardin Botanic0 de Zonas Aridas “Albert0Carballo A.”Escuela Superior de Agricultura

“Hermanos Escobar”Km. 12.5 Carr. PanamericanaCd. Juarez, Chihuahua

Distrito FederalJardin Botanic0 de La Escuela National DeMaestrosAvenida Mexico-Tacuba y Avenida de1

Maestro RuralMexico, D.F. 02801

Jardin Botanic0 de1 Instituto de BiologiaUniversidad National Autonoma de MexicoCiudad Universitaria Delegation CoyoacanMexico, D.F. 04510

Jardin Botanic0 “de La Cruz Badiano”Campus Ii. Enep Zaragoza

Apdo. Postal 9-020Mexico, D.F. 15000

DurangoJardin Botanic0 y Vivero de CactaceasDe Gomez PalacioBoulevard Gonzalez de La Vega y CalzadaLazaro CardenasCd. de Gomez Palacio, Durango

Estado De MkxicoJardin Didactic0 de CactaceasAdyacente a La Zona Arqueologica deTeotihuacanTeotihuacan, Edo. de Mexico 55831

Jardin Botanic0 de La Enep-IztacalaAv. de Los Barrios S/NSn. Juan de Los Reyes IztacalaTlalnepantla, Edo. de Mexico 54090

Jardin Botanic0 “Maximino Martinez”Universidad Autonoma de ChapingoKm. 38.5 Carr. Mexico-TexcocoChapingo, Edo. de Mexico 56230

Pinetum Maximino MartinezCentro de Genetica ForestalLomas de San Juan ChapingoApartado Postal 104Chapingo, Estado de Mexico 56230

GuanajuatoJardin BotanicoEl Charco de1 IngenioCan Te, AC.Mesones 7 1San Miguel de Allende, Gto. 37700

GuerreroJardin Botanic0 de La UniversidadAutonoma de GuerreroCiudad UniversitariaChilpancingo, Guerrero

JaliscoJardin Botanico “Jorge Victor Ellcr T.”de La Univ. Autonoma de GuadalajaraAvenida Patria No. 120 1Fraccionamiento Lomas de1 ValleZapopan, Jalisco 45 129

MichoachnOrquidario de1 CentroConvenciones de MoreliaAv. Ventura Puente S/NMorelia, Michoacan

MorelosJardin Etnobotanico de1 Inah.Matamoros 200 Col. AcapantzingoCuernavaca, More& 62440

Jardin Botanic0 de La UniversidadAutonoma de1 Estado de MorelosAv. Universidad 1001 Cal. ChamilpaCuernavaca, Morelos 622 10

Nuevo Le6nJardin Botanic0 “Efrain Hernandez X.”Facultad de Ciencias ForestalesUniversidad Autonoma de Nuevo Leon

Apdo. Postal 41Linares, Nuevo Leon 67700

PueblaJardin Botanico “ Dra Hclia Bravo “Km.25.5 Carr.Tehuacan-Hua.juapan dc

LeonZapotitlan Salinas, Pue. 75870

192

Jardin Botanic0 “Ing. RodriguezAlconedo”de La Universidad Autonoma de Puebla

Edificio Carolino4 Sur 104 Puebla, Pue. 72000

QueretaroCactario Regional y Jardin Botanic0“Hernando Sanchez Mejorada”de1 Instituto Tecnologico y de EstudiosSuperiores de Monterrey CampusQueretaro Henry Ford10 Parques Industrial. A. P. 37

Queretaro, Qro 76000

Jardin Botanico Regional de Cadel :yta“Ing. Manuel Gonzalez de Cosio”Ejido Fuentes y Pueblo NuevoCadereyta de Monks, Qro

Qintana RooJardin Botanic0 de1 CiqroKm. 320 de La Carr. Fed. 307ChetumalCancanPuerto Morelos, Q. RooApdo. Postal 886Can&m, Q. Roo 71500

Jardin Botanic0 TropicalCampo Experimental ForestalKm.40 Carr. Chetumal-CancunSn. Felipe Bacalar, Q. Roo 77930

Jardin Botanic0 TropicalSn. Felipe Bacalar”

Km. 50 de La CarreteraChetumal-Carrillo PuertoSn.Felipe Bacalar, Q. Roo

San Luis PotosiJardin Botanico de1 Institutode Invest. de Zonas DeserticasUniv. Aut. de San Luis PotosiAlvaro Obregon 64San Luis Potosi, S.L.P. 78000

SinaloaJardin Botanic0 de1 Acuario de MazatlanAv. de Los deportes No. 111Mazatlan, Sinaloa

TabascoJardin Agricola Tropical de1 CruseCentro Regional Universitario de1 SuresteApdo. Postal No. 29Teapa, Tabasco 86400

Jardin Botanic0“Parque Museo de La Venta”Av. Ruiz Cortinez S/NVillahermosa, Tabasco

TamaullpasJardin Botanico de1 InstitutoTecnologico de Cd. VictoriaBoulevard Emilio Portes Gil S/NCd. Victoria, Tamaulipasc. P. 07010

TlaxcalaJardin Botanico dc Tizatlan Tlax.Secoduci, Km. 1.5 Carr. Tlaxcala-PueTlaxcala, Tlax. 90000

VeracruzJardin Botanico de1 Ctro.RcgionalUniversitario Oriente-HuatuscoKm. 6 Carretera Huatusco-JalapaHuatusco. Ver. 94 100

Jardin Botanico “ Fco. J. Clavijero”Km. 2.5 Antigua Carr. A CoatepecXalapa, Ver. 9 1000

YucathnJardin Botanico de1 Ctro. RegionalUniversitario Peninsula de YucatanExhacienda Temozon Nor-kDesvio en El Km . 10. de La Carr. Mcrida-ProgresoMunicipio Temozon, Yucatan 97740

Jardin Botanico Regional CICY -Centro de Investigaciones Cientificas dcYucatanExhacienda XcumpichKm. 7 de La Antigua Carretera Progreso-MeridaMerida. Yucatan

193

Annex 14

Succulents of the West Indies

Compiled by Albert0 Areces-Mallea.

Table 1 Native genera

Genera marked with an asterisk (*) contain non-succulent species in the West Indies.

Total no.of species

No. ofendemic species

Total no.of species

No. ofendemic species

AGAVACEAEAgaveFurcraea

AIZOACEAECypseleaSesu viumTrianthema

AMARANTHACEAEBlu taparonLitophila

APOCYNACEAEPlumeriaRhabdadenia

ASCLEPIADACEAECynanchum *Sarcostemma

BASELLACEAEAnredera

BATACEAEBa tis

BOMBACACEAEBorn ha copsisCeibaNeobuchiaOchromaQuararibea

BORAGINACEAEArgusiaHeliotropium *

CACTACEAEAcanthocereusCereusDendrocereusDisocactusEpiphyllumEscobariaHarrisiaHylocereusLeptocereusMammillariaMelocactusOpun tiaPereskia

PilosocereusRhipsalisSelenicereusStenocereus

CARICACEAECarica

CHENOPODIACEAEHeterostachysSalicorniaSuaeda

COMPOSITAEBorrichiaIva

CRUCIFERAECa kileChamaesyce *CnidoscolusCubanthusEuphorbia *GrimmeodendronJa tropha *OmphaleaPedilan thusVictorinia

GOODENIACEAEScaevola

MORACEAEDorstenia

ORCHIDACEAEVanilla

PLUMBAGINACEAELimonium

PORTULACACEAEPortulacaTalinum

SCROPHULARIACEAEAgalinis *Bacopa *CissusHildegardiaPterocissusSuriana

334

333

231 0

00

35

001

121

1 00

32

3 10

0812

53

1526622

014

265

0

2 20

200 10 0

00

24 23

12 60022

2 113

40193

1 52

1122

4

94

153

1 228

5

00

2127

1 0

194

Table 2 Non-endemic native species

AGAVACEAEFurcraea foetida

AIZOACEAECypselea humifusaSesuvium maritimumS. portul ‘acastrumTrianthema portulacastrum

AMARANTHACEAEBlutaparon vermiculareLitophila muscoides

APOCYNACEAEPlumeria albaP. obtusaRhabdadenia biflora

ASCLEPIADACEAECynanchum angusti-foliumC. scopariumSarcostemma clausum

BASELLACEAEAnredera leptostachys

BATACEAEBatis maritima

BOMBACACEAECeiba pentandraOchroma pyramidaleQuararibea turbina ta

BORAGINACEAEArgusia gnaphalodesHeliotropium curassavicum

CACTACEAEAcanthocereus pen tagonusCereus margaritensisDisocactus ramulosusEpiphyllum phyllanthusHylocereus unda tusMammillaria mammillarisMelocactus broadwayiOpuntia caribaea0. curassavica0. dillenii0. elatior0. stricta0. wen tianaPereskia aculea taRhipsalis bacciferaSelenicereus boeckmannii

CHENOPODIACEAESalicornia bigeloviiS. perennisS. virginicaSuaeda conferta

S. fruticosaS. linearisS. tampicensisS. torreyana

COMPOSITAEBorrichia arborescensB. j?u tescensIva im brica ta

CRUCIFERAECakile lanceolata

EUPHORBIACEAEChamaesyce blodgettiC. lasiocarpaC. mesembrian themifoliaC. thymifoliaEuphorbia trichotomaOmphalea diandraPedilanthus tithymaloides

GOODENIACEAEScaevola plum ieri

MORACEAEDorstenia con trajerva

ORCHIDACEAEVanilla barbella taI/. dillonianaI/ hartiiI/. mexicanaI/. phaean thaI/ wrightii

PORTULACACEAEPortulaca ela tiorP. halimoidesP. pilosaP. quadrijidaP. rubricaulisP. teretifoliaP. urn bra ticolaTalinum .fnr ticosumT. paniculatum

SCROPHULARIACEAEAgalinis maritimaBacopa monnieri

SURIANACEAESuriana maritima

VITACEAECissus erosaC. formosaC. microcarpaC. rhom bifoliaC. trifolia taC. verticilla ta

195

Table 3 Endemic speciesDistribution1 Status2

AGAVACEAEAgave acicularisA. acklinicolaA. albescensA. anomalaA. antillarumA. bahamanaA. barbadensisA. braceanaA. brevipetalaA. brevispinaA. cacozelaA. cajalbanensisA. caribaeicolaA. dussianaA. griseaA. harrisiiA. inaguensisA. indaga torumA. in term&aA. kara ttoA. legrellianaA. longipesA. millspaughiiA. missionumA. nashiiA. papyrocarpaA. penden ta taA. shaferiA. sheuermanianaA. soboliferaA. tubulataA. underwoodiiA. van-grolaeFurcraea hexapetalaF. macrophyllaF. tu berosa

AIZOACEAECypselea ru brifloraSesuvium microphyllum

APOCYNACEAEPlumeria clusioidesP. filifoliaP. krugiiP. lanataP. montanaP. stenopetalaP. stenophyllaP. subsessilisP. trinitensisP. tu bercula ta

ASCLEPADACEAECynanchum eggersii

BOMBACACEAEBombacopsis cubensisB. emarginataNeobuchia paulinae

CACTACEAEDendrocereus nudiflorusD. undulosusDisocactus ala tusEscobaria cubensis

Harrisia brookii

Central Cuba*Acklin I (B)”East Cuba*B, CHispaniola”BBarbados*BHaiti*Haiti”BWest Cuba*LALACuba*Jamaica*BBHispaniola4’LAWest Cuba*Jamaica*BPR, VIBCentral Cuba*East Cuba”East Cuba*LACI, JWest Cuba*East Cuba*LAc, K JB, cLA, PR

PEPENENENENEPENE

ST?ST?PEPENENESTST

NENENENESTPEPENEPENEPEPEST

NENENENENENE

West Cuba* STB, CT, C, LA NE

Cuba*E. Cuba*W. Puerto Rico*E. Cuba*E. Cuba*Hispaniola*E. Cuba’”Hispaniola*Central Cuba*c, H

NESTST

NENENENENENENE

B, C, I-I NE

W. Cuba*Hispaniola*Haiti*

NENE

NE‘?

Cuba”’Hispanioloa”Jamaica*E. Cuba*Long I (B)*

NENEPEHEPE

Distribution’Hispaniola”W. Cuba”’Cuba*Jamaica*Hispaniola*Puerto Rico*W. Cuba*E. Cuba*W. Cuba”Jamaica”’H, LA, PRCentral Cuba*W. Cuba”E. Cuba*W. Cuba”’Culebra I, PR:”W. Cuba”E. Cuba*Hispaniola”W. Cuba*Puerto Rico”E. Cuba*W. Cuba*East Cuba”’Hispaniola”West Cuba”B, LA, PRC HCentral Cuba”:East Cuba”JamGca”’cCentral Cuba*East Cuba’East Cuba”B, LA, PRHispanola*West Cuba *Central Cuba*East Cuba”H;jitiS

H, LA, PRBW. PR’Florida cays*Haiti”’

Haiti*East Cuba”Jamaica”Bahamas”East Cuba”East Cuba”B, CI, CC, H, PRB, CPR, VILA, PRJamaic?‘Jamaica”Hispaniola”LA, PRJamaica*Hispaniola”Hispaniola*

Status2NENENENENEPEPENEHENENEST

NEPEPESTST

NENENENENEHEPENEHENENEHENENENENEST

NENEHEPEPE

PE?NENEHEHE

PE‘.’PE?NENEPENENENENENENENEPENENENENEPENE

H. divarica taH. earleiH. eriophoraH. gracilisH. nashiiH. portoricensisH. taetraH. tayloriHylocereus cu bensisH. tria ngula risH. trigonusLeptocereus arboreusL. assurgensL. carina tusL. ekmaniiL. grantianusL. leoniiL. maxoniiL. panicula tusL. prostra tusL. quadricosta tusL. santamarinaeL. scopulophilusL. sylvestrisL. weingartianusL. wrigh tiiMammillaria nivosaM. proliferaMelocactus actinacanthusM. acunaiM. communisM. guitartiM. harlowiiM. holguinensisM. in tortusM. lemaireiM. matanzanusM. perezassoiM. radocziiOpuntia acaulis0. an tillana0. bahamana0. borinquensis0. corallicola0. ekmanii0. falcata0. hystrix0. jamaicensis0. lucayana0. macracantha0. militaris0. millspaughii0. monilifonnis0. nashii0. repens0. ru bescens0. sanguinea0. spinosissima0. taylori0. triacan tha0. tunaPereskia marcanoiP. portulacifolia

196

P. quisqueyanaP. zinniifloraPilosocereus bahamensisP. brooksianusP. millspaughiiP. nobilisP. polygonusP. robiniiP. royeniiP. swartziiSelenicereus brevispinusS. grandiflorusS. innesiiS. urbanianusStenocereus hystrix

CARICACEAECarica jamaicensis

CHENOPODIACEAEHeterostachys ritteriana

COMPOSITAEBorrichia cubana West Cuba”Iva cheiranthifolia B, CI, C

EUPHORBIACEAEChamaesyce abbrevia taC. brittoniiC. cayensisC. cowelliiC. hepaticaC. lecheoidesC. vaginula taC. wilsoniiCnidoscolus bella torC. fragransC. matosiiC. quinquelobatusC. range1Cubanthus brittoniiC. linearifoliusC. umbellif&nisEuphorbia ala taE. cassythoidesE. cubensisE. defolia taE. dussiiE. eggersiiE. gymnonotaE. helenaeE. longinsulicolaE. marchiiE. muniziiE. petiolarisE. podocarpifoliaE. puniceaGrimmeodendron eglandul sumG. jamaicenseJatropha angustifoliaJ. divarica taJ. hemandiifoliaJ. in tegerrimaJ. minorJ. tupifoliaOmphalea commutata0. ekmanii0. hypoleuca0. triandra0. trichotoma

Hispaniola”Cuba*BEast Cuba*B, cLA (Windward I.)c, HWC, F caysLA, PRCI, JCuba*CI, C, JSt. Vincent’”

c, HC, J, PR

Jamaica*

Hispaniola*

BBahamas*BH, PRH, NavassaBBBWest Cuba*West Cuba*East Cuba”West Cuba*West Cuba*Central Cuba*Cuba*c, HJamaica*B, CI, C, HWest Cuba*Hispaniola*Mart., St. Lucia”Hispaniola*BEast CubaLong I. (B)”East Cuba*East Cuba”B, GA, LAEast Cuba*Jamaica*B, C, HJamaica*Cuba*Jamaica*H, J‘?, PRCuba*East Cuba*Cuba*Hispaniola*Hispaniola*West Cuba*Jamaica*West Cuba*

HEST

NENENENENEST

NENEPENEHENENE

NE

NE

HENE

NENENENENENENENENEHEPEHENENENENEPENENESTPENENENEPENENENENENENEPENENENENENENENENENENENE

Pedilanthus bahamensisVictorinia acrandraI/: regina

MORACEAEDorstenia caimitensisD. cordifoliaD. crassipesD. cren ula taD. domingensisD. ekmaniiD. erythrandraD. fa wcettiiD. flagelliferaD. hotteanaD. howardiiD. jamaicensisD. laneiD. marginataD. multisquameaD. nipensisD. nummulariaD. peltataD. petraeaD. rocanaD. roigiiD. tricolorD. tuberosa

ORCHIDACEAEVanilla claviculataV. correlliiI/: pleeiI/: poitaeiK rubraV savanarum

PLUMBAGINACEAELimonium bahamense

PORTULACACEAEPortulaca brevijoliaP. caulerpoidesP. minutaP. tu bercula taTalinum domingense

STERCULIACEAEHildegardia c-11 bensis

VITACEAECissus carnifc~liuC. corullicolaC. dichroaC’. dissectaC. fuertessiiC. gonavensisC. grisebachiiC. haitiensisC. hotteanaC. in termediaC. micran thaC. morn icolaC. n ipensisC. obovataC. parcifk,raC. picardaeC. ru brinerviuC. rupicolaC. subaveniuC. torreanaC. tuberculataPterocissus m ira bilis

Distribution1B

status

Hispaniola’”East Cuba”:

NEPEPE

Hispaniola:”Hispaniola”’East Cuba *East Cuba”Hispaniola”’Central CubaEast Cub,1 ‘:cJamaica”’Hispaniola Ii’Hispaniola+’East Cuba *Jamaica”’Central CubaHispaniola’”Hispaniola”’East Cuba”:Central CubaHispaniokFEast Cuba :‘:Central Cubawest Cuba”~East Cuba:‘:East Cuba’!:

,<T’?i)“ip

PE‘I“T”?

J\jEI’E

4E’T

(E: ,7’?s I?NENEST

ST’.’ST’!NEPESTNENENENE

ST?

CI, GABahamas”’Mart. (LA)‘”H, PRHaiti”’East Cuba”:

PE

C, HaitiW PR”’BCI, E, CHispaniola”’

NENENEPEPE

East Cuba”’ ST

Haiti3:

East Cuba”’w e s t C u b a ” :

Hispaniola”’Hispaniola”’Hispaniola:”C, HHaiti:%

Hispaniola”B, GAHispaniola:’Hispaniola:”East Cub?cGA, LAHispaniola*’Hispaniola:i’Hispaniola’i’west Cuba”:East Cuba ‘:CUba 9:

B, GAHispaniola”’

PEST

NE:NENEPENE

E-I E’.’NENENEPENENEPENENENENENEN E;:NE

197

‘An asterisk (*) indicates restricted distribution; abbreviated location indicates regional or widespread distribution. B Bahamas, C Cuba, CI CaymanIslands, E. East, F Florida, GA Greater Antilles, H Hispaniola, J Jamaica, LA Lesser Antilles, Mart. Martinique, N. North, PR Puerto Rico,S. South, St. Saint, VI Virgin Islands, W. West

“Conservation categories:NE ‘Not endangered’ stands for those taxa commonly represented by a relatively large number of individuals often distributed in geographicallywidespread populations, which are not facing any inminent threat at present. There are 155 endemics in this category.

PE ‘Potentially endangered’ is mostly assigned to those highly vulnerable species represented by one or few populations, and few individuals therein,that are not threatened at present but with very little degrading effort can be wiped out of the wild. Forty-six endemic species are listed as such.

ST ‘Somewhat threatened’ applies to those taxa whose strained wild populations have had a more or less significant detrimental impact, that ifmaintained could lead to the extinction of the species. The plants under this category are expected to recover relatively easily if the strain ceases andadequate sound management plans for protection are applied. There are 28 species given this status.

HE ‘Highly endangered to nearly extinct’ is the hottest category; most of the species listed here are unsalvagable unless very urgent actions arc takenimmediately to preserve them for posterity. Unfortunately, for Opuntia borinquensis, Cnidoscolus fragrans, and C. quinquelobatus it seems that nothingcan possibly be done at this moment. Fourteen species in total fall into this conservation category.

198

Annex15

Brazilian cacti

Compiled by Nigel P. Taylor.

The new IUCN categories of threat (see Annex 16 for definitions) are given below for the Cactaceae of eastern Brazil and the Brazilian Atlantic Forest,geographically defined as in the regional account of South America. The order of taxa is systematic. Endemics are indicated by an asterisk (:“).Introduced, naturalised, and most hybrid taxa are omitted.

Taxon Conservation information

PERESKIOIDEAEPereskia aculea taP. grandifolia ssp. grandqolia *

ssp. violacea *P. ba hiensis *P. stenantha *P. aureiflora *

OPUNTIOIDEAQuiabentia zehntneri *Tacinga braunii *T. finalis*Opuntia werneri *0. palmadora *0. saxatilis ssp. saxatilis *

ssp. estevesi?

0. inamoena *0. x quipa *0. monacanthaBrasiliopuntia brasiliensis

CACTOIDEAEPseudoacathocereus brasiliensis *Harrisia adscendens *Selenicereus setaceusEpiphyllum phyllanthusLepismium crucif~x-meL. houlletianum *L. warmingianumL. lumbricoidesRhipsalis pulchra *R. puniceodiscus *R. hoelleri *R. neves-armondii *R. dissimilis *R. trigona *R. jloccosa ssp. floccosa

ssp. pulvinigera *R. paradoxa

ssp. septentrionalis *ssp. paradoxa *

R. pacheco-leonis ssp. catenulata *ssp. pacheco-leonis *

R. pen tap tera *R. sulca ta *

Low Risk.Data Deficient, but perhaps Low Risk, since it has been taken into cultivation (as a hcdgc plant) in the

region where it is native.Data Deficient, ibid.Low Risk; it has been taken into cultivation (as a hedge plant) in the region whcrc it is native.Low Risk; it has been taken into cultivation (as a hedge plant) in the region where it is native.Vulnerable; this plant is rare except in north-eastern Minas Gerais (Rio Jequitinhonha valley).

Low Risk and perhaps locally increasing by vegetative propagation when its habitat is disturbed.Vulnerable.Low Risk at present, but habitat destruction continuing.Vulnerable; its habitat at Rui Barbosa (BA) is being destroyed by mining operations.Low Risk.Low risk at present, but found only on limestone and therefore potentially at risk from quarrying

activities in the long term.Data Deficient, but probably vulnerable, since it is presently known only from one location and could bc

affected by quarrying in the future.Low Risk.Low Risk.Low Risk.Data Deficient; much of its habitat has been destroyed in Brazil, but it dots occur in some Parqucs

Nacionais and has a wide distribution in the southern Neotropics.

Vulnerable.Low Risk.Low Risk.Low Risk.Low Risk.Low Risk.Low Risk.Low Risk.Data Deficient; possibly Conservation Dependent, since it is found within various protected arcas.Low Risk.Data Deficient, but to date collected only once.Low Risk, but needs to be monitored.Data deficient; included within at least one protected arca in the southern part of its range.Low Risk, but needs to be monitored.Low Risk.Low Risk.

Data deficient, but potentially endangered in view of continuing habitat destruction.Low Risk.Data deficient, but known from only two localities.Vulnerable; known only from the region of Cabo Frio (RJ), where considerable development for

tourism is taking place.Extinct in the Wild.Data Deficient, but currently known from only a single collection.

199

R. russellii *

R. elliptica *R. pachyptera *R. oblonga *R. crispa ta *

R. cereoides *

R. densiareola ta *R. baccifera ssp. baccifera

ssp. hileiabaiana *R. lindbergiana *R. teres*R. grandiflora *R. mesembryanthemoides *R. juengeri *R. cereusculaR. pilocarpa *

R. burchellii *

R. ewaldiana *R. campos-portoana *R. clavata *Hatiora salicomioides *H. herminiae *

H. epiphylloides ssp. epiphylloides *

ssp. bradei *H. gaertneri *H. rosea *Schlumbergera kau tskyi *S. russelliana *S. truncata *S. orssichiana *S. microsphaerica *S. opuntioides *Brasilicereus phaeacanthus *B. markgrafii *Cereus euchlorusC. mirabella *C. albicaulis *C. fernambucensis ssp. femambucensis *

ss p. sericifer *C. insular-is *C. jamacaru ssp. jamacaru *

ssp. calcirupicolaC. hildmannianusCipocereus laniflorus* (sp. nov. ined.)

C. crassisepalus *C. bradei”C. minensis ssp. minensi?

ssp. pleurocarpus *

C. pusilliflorus *

Stephanocereus leucostele *S. luetzelburgii *Arrojadoa bahiensis *

Low Risk; however, its habitat has contracted markedly especially in southern Bahia, where the mostrecent collection was made in 1971. Elsewhere, it is found within some reasonably secure protectedareas.

Low Risk taking its total range into account.Low Risk.Low Risk taking its total range into account.Data Deficient. Apparently wide-ranging but rarely observed and potentially vulnerable despite its

range.Vulnerable; known from only two localities, the southern one close to the city of Rio de Janeiro and

potentially affected by coastal tourism.Data Deficient.Low Risk.Vulnerable.Not threatened.Low Risk.Low Risk, but needs to be monitored.Vulnerable; situated close to the city of Rio de Janeiro.Data Deficient.Low Risk.Vulnerable or Conservation Dependent; found only in virgin forest, very little of which remains (some

of it possibly in National Parks or other reserves).Vulnerable; its type locality (one of only five known for the species) is now within the city of Sao Paula

and has been heavily polluted in recent times.Data Deficient.Low Risk.Low Risk or possibly Conservation Dependent, since included in various protected areas.Low Risk.Vulnerable and probably dependent on the maintenance of its habitat in the Parque Estadual Campos

do Jordao to avoid it becoming endangered or critical.Endangered; known only from the type locality, now situated inside a National Park, but not known to

have been seen or collected in recent times.Vulnerable; known only from a single habitat, only part of which is included inside a protected area.Data Deficient.Data Deficient, but relatively wide-ranging and possibly Low Risk.Data deficient, but potentially vulnerable or endangered.Conservation Dependent?Conservation Dependent?Data Deficient; possibly included within a protected area, but range poorly understood.Data deficient; included within some protected areas.Data deficient; included within some protected areas.Low Risk.Vulnerable.Low Risk.Vulnerable.Low Risk, but habitat is declining.Low Risk.Low Risk.Low Risk.Low Risk.Low Risk, but habitat is declining in some places due to quarrying of limestone.Low Risk.Data Deficient. Presently known from only a single, small population (< 50 mature individuals), but

further field investigations are needed to determine range and abundance.Vulnerable, from habitat destruction by charcoal producers.Vulnerable; of very limited range and not included within any protected area.Low risk.Vulnerable from excessive burning of its habitat, which is partly within the Parquc National da Serra do

Cipo.Critically Endangered on present knowledge and urgently in need of further field studies to determine

if its range is more extensive.Low Risk at present, but its habitat is continuing to decline and its status needs to be monitored.Low Risk.Vulnerable: known from only four populations (one within the Parque National da Chapada

Diamantina) and at risk from collection of plants and seeds. However, many individuals arc protectedby the plant’s preference for steep slopes and cliffs inaccessible to the collector.

200

A. dinae ssp. dinae*ssp. eriocaulis *

A. penicilla ta *A. rhodantha *Pilosocereus tuberculatus *P. gounellei ssp. gounellei *

ssp. zehntneri *P. ca tingicola ssp. ca tingicola *

ssp. salvadorensis *P. sp. nov. ined. [Pedra Azul, MG]“’

P. arra bidae *P. brasiliensis ssp. brasiliensis *

ssp. ruschianus *P. flavipulvina tus *

P. pentaedrophorus ssp. pentaedrophorus”ssp. robustus *

P. glaucochrous *P. jIoccosus ssp. jIoccosus *

ssp. quadricostatus *P. albisummus *P. flexibilispinus *P. ulei”

P. fulvilanatus ssp. fulvilanatus *ssp. rosae *

P. pachycladus ssp. pachycladus *ssp. pemambucoensis *

P. magnificus *P. machrisiiP. aurisetus ssp. aurisetus *

ssp. aurilanatus *

P. aureispinus *P. vila boensis *P. multicosta tus *P. piauhyensis *P. chrysostele *P. diersianus *P. densiareola tus *Micranthocereus violacijlorus *

M. albicephalus *M. purpureus *M. auriazureus *M. streckeri *M. polyanthus*M. jlaviflorus *M. dolichospermaticus *

M. estevesii *Coleocephalocereus buxbaumianus *

C. fluminensis ssp. decumbens *ssp. fluminensis *

C. pluricosta tus *C. goebelianus *C. aureus *C. purpureus *

Melocactus oreas ssp. oreas *ssp. cremnophilus *

M. emestii ssp. ernestit?ssp. longicarpus *

Vulnerable from habitat destruction by charcoal producers.Vulnerable from habitat destruction by charcoal producers. Needs regular monitoring if it is not to

become endangered.Low Risk, but its habitat continues to decline.Low Risk, but its habitat continues to decline.Low Risk, but its habitat continues to decline.Low Risk.Low Risk, but since many populations are found on limestone it may decline due to quarrying activities.Low Risk, but its habitat continues to decline and its status needs to be monitor-cd.Low Risk, ibid.Data Deficient, but possibly Vulnerable, since known populations inhabit forest vcgctation which could

be cleared for agriculture or charcoal production.Low Risk.Data Deficient, but has apparently declined markedly in the southern part of its range (Rio dc Janeiro).

Low Risk.Data Deficient, but has probably declined in the southern part of its range due to forest clearance

(Piaui).Low Risk, but its habitat continues to decline.Low Risk, but its habitat continues to decline.Low Risk, but its habitat continues to decline.Low Risk, but its habitat on limestone outcrops is at risk from quarrying operations.Vulnerable from clearance of its caatinga habitat.Data Deficient, but known from only a single site.Data Deficient, ibid.Vulnerable; known only from a very limited area which continues to be devclopcd for tourism (C’abo

Frio, RJ).Vulnerable from habitat disturbance and destruction for charcoal production.Data Deficient, but possibly vulnerable from habitat disturbance caused by local industry.Low Risk, but caatinga habitat continues to decline (camp0 rupestre habitats less threatcncd).Low Risk, but habitat continues to decline.Low Risk, but range limited.Low Risk.Low Risk.Vulnerable; known only from a small region without protected areas, the surrounding habitats at risk

from charcoal production activities.Data Deficient. Range may be much greater than presently known.Data Deficient; known from only two sites.Low Risk, but needs to be monitored in view of its restricted distribution.Data Deficient.Data Deficient.Data Deficient, but known from only a single site.Low Risk, but habitat liable to decline through limestone quarrying.Vulnerable and liable to become Endangered if charcoal producers move into the restricted arca whcrc

it occurs. It needs regular monitoring.Vulnerable.Low Risk.Vulnerable, due to its restricted distribution.Data Deficient, but possibly Endangered or Vulnerable, due to its very restricted distribution.Vulnerable, due to its restricted distribution.Low Risk.Data Deficient, but probably vulnerable from destructive collection of seeds and from potential

quarrying of limestone.Data Deficient, ibid.Low Risk in the eastern sector of its range, but Endangered to the west, where mining operations have

destroyed its habitat.Data Deficient, but vulnerable at its type locality from urban expansion.Low Risk.Low Risk, since its habitats are mostly inaccessible, steep slopes.Low Risk at present, but its habitat continues to be lost.Low Risk.Data Deficient, but perhaps vulnerable, since the known habitat is close to a road and may lx visited by

collectors for plants and seed.Low Risk at present, but further modification of its habitats is likely and it requires monitoring.Low Risk.Low Risk.Low Risk at present, but habitats are being disturbed in certain parts of its range; needs to bc

monitored.

201

M. ba hiensis * ssp. bahiensis *ssp. amethystinus *

M. conoideus *

M. deinacanthus *

M. levites ta tus *M. azureus ssp. azureus”

ssp. ferreophilus *M. pachyacanthus ssp. pachyacanthus *

ssp. viridis *M. salvadorensis *M. zehntneri *M. lanssensianus *M. glaucescens *

M. concinnus *M. paucispinus *

M. violaceus ssp. violaceu?ssp. ritteri *ssp. margaritaceus *

Leocereus bahiensis *Facheiroa ulei *F. cephaliomelana ssp. cephaliomelana *

ssp. estevesii *F. squamosa *Espostoopsis dybowskii *

Arthrocereus melanurus ssp. melanurus *ssp. magnus [ssp. nov. ined.]*ssp. odor-us *

A. rondonianus *A. glaziovii *Discocactus zehntneri ssp. zehntneri *

ssp. boomianus *D. ba hiensis *

D. heptacanthusD. platen tiformis *D. pseudoinsignis *D. horstii *Uebelmannia buiningii *

U. gummifera *

U. pectinifera ssp. pectinifera *ssp. flavispina *ssp. horrida *

Low Risk.Low Risk.Critically Endangered. Regrettably, survival of M. conoideus in the wild is severely threatened by

extraction of the quartz gravel in which it grows, and through commercial collection for the Europeanhorticultural market. Since June 1992 it has been listed on Appendix I of CITES

Vulnerable or Endangered, depending on its status at the locality in Municipio Born Jesus da Lapa,which has not been visited. In view of its rarity and vulnerability to commercial exploitation it has beenplaced on Appendix I of CITES since June 1992.

Low Risk at present, but in need of monitoring in view of the potential threat from limestone quarrying.Endangered by actual or potential habitat destruction at its few known localities.Vulnerable from limestone quarrying.Endangered by actual or potential habitat destruction at its few known localities.Critically Endangered by habitat destruction at its known localities.Low Risk at present, but its habitat continues to disappear.Low Risk.Data Deficient.Endangered. In view of its rarity and the threats from commercial collection M. glaucescens has been

placed on Appendix I of CITES since 1992.Low Risk, but some populations affected by vegetation burning (campos rupestres).Endangered. In view of its restricted distribution, rarity and desirability to collectors this species has

been listed in Appendix I of CITES since 1992.Vulnerable from habitat modification by the tourist industry.Vulnerable. Known from only two locations, both near to towns.Vulnerable from habitat modification by the expanding tourist industry.Low Risk.Data Deficient.Data Deficient, but restricted to limestone which may be quarried in future.Vulnerable, with the potential for future limestone quarrying at its only known site.Low Risk.Vulnerable; both populations of limited extent, the northern one affected by clearance of the caatinga

in some places.Vulnerable.Vulnerable due to its very limited range, but located within a protected area.Vulnerable.Vulnerable.Vulnerable; many of its former habitats have been eliminated through the mining of iron oreVulnerable; part of its former habitat has been submerged beneath the Represa de Sobradinho.Vulnerable; subject to collection, at least one of its habitats being accessible from a main road.Data Deficient, but part of its range was eliminated by inundation from the Represa de Sobradinho and

some of its few sites are accessible by road and may be visited by collectors.Low Risk, but in need of monitoring, since the cerrado habitats are undergoing much change.Vulnerable, since most populations are small.Vulnerable; range restricted.Endangered; range very restricted and heavily impacted by collectors in the recent past.Critically Endangered. According to Braun & Esteves Pereira this species is on the verge of extinction

and is scarcely if at all in cultivation, where it has proved very difficult. Affected by collection of plantsand seed.

Endangered. Range very limited and habitat affected by charcoal production and collection of plantsand seed.

Vulnerable, from collection of plants and seed.Vulnerable, from collection of plants and seed.Vulnerable; known from only a single locality.

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Annex 16

The IUCN Red List Categories

Pre-I994 categories (IUCN 1980)

Extinct (Ex). Taxa which are no longer known to exist in the wild after repeated searches of their type localities and otherknown or likely places.

Endangered (E). Taxa in danger of extinction and whose survival is unlikely if the causal factors continue operating. Includedare taxa whose numbers have been reduced to a critical level or whose habitats have been so drastically reduced that they aredeemed in immediate danger of extinction.

Vulnerable (V). Taxa believed likely to move into the Endangered category in the near future if the causal factors continueoperating. Included are taxa of which most or all the populations are decreasing because of over-exploitation, extensivedestruction of habitat or other environmental disturbance; taxa with populations that have been seriously depleted and whoseultimate security is not yet assured; and taxa with populations that are still abundant but are under threat from serious adversefactors throughout their range.

Rare (R). Taxa with small world populations that are not at present Endangered or Vulnerable, but are at risk. These taxa areusually localized within restricted geographical areas or habitats or are thinly scattered over a more extensive range.

Indeterminate (I). Taxa known to be Extinct, Endangered, Vulnerable or Rare, but where there is not enough information tosay which of the four categories is appropriate.

Insufficiently Known (K). Taxa that are suspected but not definitely known to belong to any of the above categories because ofa lack of information.

Not threatened (nt). Taxa that are not in any of the above categories.

No Information or Unknown (?). Taxa for which there is no information.

Note: In addition to the categories above, occasionally “hybrid” categories are used, such as Ex/E (probably Extinct) or E/V(near Endangered).

The IUCN Red List Categories

Prepared by the IUCN Species Survival CommissionAs approved by the 40th Meeting of the IUCN Council, Gland, Switzerland, 30 November 1994

The categories

Extinct (EX)A taxon is Extinct when there is no reasonable doubt that the last individual has died.

Extinct in the Wild (EW)A taxon is Extinct in the Wild when it is known only to survive in cultivation, in captivity or as a naturalised population (orpopulations) well outside the past range. A taxon is presumed extinct in the wild when exhaustive surveys in known and/orexpected habitat, at appropriate times (diurnal, seasonal, annual), throughout its historic range have failed to record anindividual. Surveys should be over a time frame appropriate to the taxon’s life cycle and life form.

203

Critically Endangered (CR)A taxon is Critically Endangered when it is facing an extremely high risk of extinction in the wild in the immediate future, asdefined by any of the criteria (A to E) below.

Endangered (EN)A taxon is Endangered when it is not Critically Endangered but is facing a very high risk of extinction in the wild in the nearfuture, as defined by any of the criteria (A to E) below.

Vulnerable (VU)A taxon is Vulnerable when it is not Critically Endangered or Endangered but is facing a high risk of extinction in the wild inthe medium-term future, as defined by any of the criteria (A to D) below.

Lower Risk (LR)A taxon is Lower Risk when it has been evaluated, does not satisfy the criteria for any of the categories Critically Endangered,Endangered or Vulnerable. Taxa included in the Lower Risk category can be separated into three subcategories:

Conservation Dependent (cd). Taxa which are the focus of a continuing taxon-specific or habitat-specific conservationprogramme targeted towards the taxon in question, the cessation of which would result in the taxon qualifying for one of thethreatened categories above within a period of five years.

Near Threatened (nt). Taxa which do not qualify for Conservation Dependent, but which are close to qualifying for Vulnerable.

Least Concern (1~). Taxa which do not qualify for Conservation Dependent or Near Threatened.

Data Deficient (DD)A taxon is Data Deficient when there is inadequate information to make a direct, or indirect, assessment of its risk of extinctionbased on its distribution and/or population status. A taxon in this category may be well studied, and its biology well known, butappropriate data on abundance and/or distribution is lacking. Data Deficient is therefore not a category of threat or LowerRisk. Listing of taxa in this category indicates that more information is required and acknowledges the possibility that futureresearch will show that threatened classification is appropriate. It is important to make positive use of whatever data areavailable. In many cases great care should be exercised in choosing between DD and threatened status. If the range of a taxonis suspected to be relatively circumscribed, if a considerable period of time has elapsed since the last record of the taxon,threatened status may well be justified.

Not Evaluated (NE)A taxon is Not Evaluated when it is has not yet been assessed against the criteria.

The Criteria for Critically Endangered, Endanpered and Vulnerable

CRITICALLY ENDANGERED (CR)A taxon is Critically Endangered when it is facing an extremely high risk of extinction in the wild in the immediate future, asdefined by any of the following criteria (A to E):

A) Population reduction in the form of either of the following:

1) An observed, estimated, inferred or suspected reduction of at least 80% over the last 10 years or three generations,whichever is the longer, based on (and specifying) any of the following:a) direct observationb) an index of abundance appropriate for the taxonc) a decline in area of occupancy, extent of occurrence and/or quality of habitatd) actual or potential levels of exploitatione) the effects of introduced taxa, hybridisation, pathogens, pollutants, competitors or parasites.

204

B)

2) A reduction of at least 80%, projected or suspected to be met within the next 10 years or three generations,

whichever is the longer, based on (and specifying) any of (b), (c), (d) or (e) above.

Extent of occurrence estimated to be less thanestimates indicating any two of the following:

100 km2 or area of occupancy estimated to be less than 10 kmz, and

1) Severely fragmented or known to exist at only a single location.

2) Continuing decline, observed, inferred or projected, in any of the following:a) extent of occurrenceb) area of occupancyc) area, extent and/or quality of habitatd) number of locations or subpopulationse) number of mature individuals.

3) Extreme fluctuations in any of the following:a) extent of occurrenceb) area of occupancyc) number of locations or subpopulationsd) number of mature individuals.

C) Population estimated to number less than 250 mature individuals and either:

1) An estimated continuing decline of at least 25% within three years or one generation, whichever is longer or

2) A continuing decline, observed, projected, or inferred, in numbers of mature individuals and population structure inthe form of either:a) severely fragmented (i.e. no subpopulation estimated to contain more than 50 mature individuals)b) all individuals are in a single subpopulation.

D) Population estimated to number less than 50 mature individuals.

El Quantitative analysis showing the probability of extinction in the wild is at least 50% within 10 years or threegenerations, whichever is the longer.

ENDANGERED (EN)A taxon is Endangered when it is not Critically Endangered but is facing a very high risk of extinction in the wild in the nearfuture, as defined by any of the following criteria (A to E):



2) A reduction of at least 50%, projected or suspected to be met within the next 10 years or three generations,whichever is the longer, based on (and specifying) any of (b), (c), (d), or (e) above.

B) Extent of occurrence estimated to be less than 5000 km2 or area of occupancy estimated to be less than 500 km2, andestimates indicating any two of the following:

1) Severely fragmented or known to exist at no more than five locations.

205

2) Continuing decline, inferred, observed or projected, in any of the following:

a) extent of occurrenceb) area of occupancyc) area, extent and/or quality of habitatd) number of locations or subpopulationse) number of mature individuals.

3) Extreme fluctuations in any of the following:a) extent of occurrenceb) area of occupancyc) number of locations or subpopulationsd) number of mature individuals.

C) Population estimated to number less than 2500 mature individuals and either:

1) An estimated continuing decline of at least 20% within five years or two generations, whichever is longer, or

2) A continuing decline, observed, projected, or inferred, in numbers of mature individuals and population structure inthe form of either:a) severely fragmented (i.e. no subpopulation estimated to contain more than 250 mature individuals)b) all individuals are in a single subpopulation.

D) Population estimated to number less than 250 mature individuals.

E) Quantitative analysis showing the probability of extinction in the wild is at least 20% within 20 years or fivegenerations, whichever is the longer.

VULNERABLE (Vu)A taxon is Vulnerable when it is not Critically Endangered or Endangered but is facing a high risk of extinction in the wildin the medium-term future, as defined by any of the following criteria (A to E):



2) A reduction of at least 20%, projected or suspected to be met within the next ten years or three generations,

whichever is the longer, based on (and specifying) any of (b), (c), (d) or (e) above.

B) Extent of occurrence estimated to be less than 20,000 km2 or area of occupancy estimated to be less than 2000 km’, andestimates indicating any two of the following:1) Severely fragmented or known to exist at no more than ten locations.

2) Continuing decline, inferred, observed or projected, in any of the following:a) extent of occurrenceb) area of occupancyc) area, extent and/or quality of habitatd) number of locations or subpopulationse) number of mature individuals

206

3) Extreme fluctuations in any of the following:a) extent of occurrenceb) area of occupancyc) number of locations or subpopulationsd) number of mature individuals

C) Population estimated to number less than 10,000 mature individuals and either:

1) An estimated continuing decline of at least 10% within 10 years or three generations, whichever is longer, or

2) A continuing decline, observed, projected, or inferred, in numbers of mature individuals and population structure inthe form of either:a) severely fragmented (i.e. no subpopulation estimated to contain more than 1000 mature individuals)b) all individuals are in a single subpopulation

D) Population very small or restricted in the form of either of the following:

1) Population estimated to number less than 1000 mature individuals.

2) Population is characterised by an acute restriction in its area of occupancy (typically less than 100 km?) or in thenumber of locations (typically less than five). Such a taxon would thus be prone to the effects of human activities (orstochastic events whose impact is increased by human activities) within a very short period of time in anunforeseeable future, and is thus capable of becoming Critically Endangered or even Extinct in a very short period.

E) Quantitative analysis showing the probability of extinction in the wild is at least 10% within 100 years.

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Annex 17

IUCN/SSC Cactus and Succulent Plant SpecialistGroup members and contributing authors

* Author, + Contributor. The first number listed below the address is telephone; the second, when listed, is fax.

Dr Bryan R. ADAMSNational Herbarium and Botanic GardenP.O. Box 8100CausewayHarareZIMBABWE2630-725313

Dr Focke ALBERS *Westfaelische Wilhelms-UniversitaetInstitut fur BotanikSchlossgarten 3Munster48149 [email protected]

Dr Edward ANDERSON (Chair) *Desert Botanical Garden1201 N. Galvin ParkwayPhoenix, Arizona85008 [email protected]

Prof Albert0 E. ARECES-MALLEA *New York Botanical Garden,Institute of Systematic BotanyBronx, New York10458-5126 USA1-718-817-8677l-718-562-6780aareces<@nybg.org

SPECIALIST GROUP MEMBERS

Mr Salvador ARIASInstituto de Biologia,Universidad National AutonomaApdo. Postal 70-614Mexico, D.F.045 10 MEXICO52-5-622-904852-5-616-2326

Dr Gerald S. BARADCactus and Succulent Society of America521 Sergeantsville RoadFlemington, New Jersey08822 [email protected]

Dr Wilhelm BARTHLOTTBotanical Institute, University of BonnMeckenheimer Allee 170Bonn53115 GERMANY49-228-732-52649-228-733-120

Dr Pierre BRAUNIm Fussta137Kerpen50171 [email protected]

Mr Hans BRITSCHP.O. Box 2018Vista, California92085 USAl-619-726-17101-619-726-1799

208

Mrs Susan CARTER-HOLMES *Royal Botanic Gardens, KewRichmond, SurreyTW9 3AB UNITED KINGDOM44-181-332-524344-181-332-5278

Dr Andrea CATTABRIGA10s Generic Reserve Collection Sectionc/o Fotolito BononiaVia Irnerio 42Bologna40126 ITALY39-51-625-573739-5 [email protected]

Dr Miguel CHAZAROFacultad de Geografia, Universidad de GuadalajaraApartardo Postal 30Zapopan, Jalisco45101 MEXICO52-3-624-136352-3-653-5058

Dr Richard FELGERDrylands Institute2509 North Campbell Ave # 176Tucson, Arizona85719 USAl-602-321-1825l-602-321-1872

Mr W.A. and Betty FITZ MAURICE *Can Te, A.C.Hermanos Infante 225San Luis Potosi, SLP78250 MEXICO52-48-131-32352-48-131-323

Prof Abisai GARCIA MENDOZA +Universidad National Autonoma de MexicoJardin Botanico, UNAMApdo. Postal 70-614Delegation CoyoacanMexico D.F.045 10 MEXICO52-5-622-897552-5-616-2326

Mr Michael G. GILBERT +Natural History Museum, Dept. of BotanyCromwell RoadLondonSW7 5BD UNITED KINGDOM44-171-938-941344-171-938-9260

Mr Charles GLASS +Can Te, A.C.Mesones 71San Miguel de Allende, Guanajuato37700 MEXICO52-415-2299052-415-24015

Mr Lambertus E. GROENHeelsumseweg 49GR Bennekom6721 [email protected]

Mr Chuck HANSONArid Lands Greenhouses3560 W Bilby RoadTuscan, Arizona85746 USAl-602-833-9404l-602-883-8874

Dr Bruce HARGREAVESNational Museum, BotswanaP.O. Box 10098The Village, GaboroneBOTSWANA267-373-860267-302-797

Dr Hector HERNANDEZInst. de Biologia, UNAMCiudad UniversitariaApartado Postal 70-233Mexico D.F.045 10 [email protected]

209

Mr Craig HILTON-TAYLOR *National Botanical InstitutePrivate Bag X7Claremont7735 SOUTH [email protected]

Dr David HUNT83 Church StreetMilborne PortDorsetSherborneDT9 5DJ UNITED KINGDOM44-1963-250-89944-1963-250-999

Dr Gary J. JAMES214 Eden Ave.Costa Mesa, California92627 USAl-714-642-2092

Prof Dr Norbert JURGENSBotanical Institute, University ofGyrhofstr. 15- .Cologne50932 [email protected]

Dr Roberto KIESLING +Instituto de Botanica DarwinionCasilla Postal 221642 San IsidroProv. Buenos AiresARGENTINA54-l-747-474854-l-747-4748

Mr Michael J. KIMBERLEYP.O. Box MP 50Mount PleasantZIMBABWE263-4-726-041

Dr Robert KRAUS *L. Anzengruber Str. 119Dorfen84405 GERMANY

(OrIInstitut fur Systematische Botanik der UniversitatMenzingerstr. 6780638 MiinchenGERMANY

Dr Beat Ernst LEUENBERGERBotanischer Garten & Botanisches MuseumBerlin-DahlemBerlin14191 GERMANY49-30-8300-611749-30-8300-6186leu@fub46.zedat.fu-berlin.de

Prof Alejandro N. LIMAMiami Dade Community College627 SW 27th AvenueMiami, Florida33135-2297 [email protected]

Mr Gary W. LYONS +2606 West Clark AvenueBurbank, California91505 USA1-818-843-5172l-818-843-5172

Dr Bruce MACBRYDEOffice of Scientific AuthorityUS Fish & Wildlife Service725 Arlington. Sq. Bldg.Washington, DC20240 USAl-703-358-1708l-703-358-2276

Mr Detlev METZINGIFOE, Abt. Geobotanik & Naturschutz, FBUniversit BremenBremen28344 GERMANY49-421-218-27349-421-218-705

Mrs Lindah MHLANGAUniversity Lake Kariba Research StationP.O. Box 48KaribaZIMBABWE263-61-22312

210

Dr Gary NABHAN +Arizona Sonoran Desert Museum3021 North Kinney RoadTucson, Arizona85743-8918 USAl-602-883-2500l-602-883-2500

Prof Leonard E. NEWTON *Kenyatta UniversityDepartment of BotanyP.O. Box 43844NairobiKENYA254-2-810-901*2

MS Sara Felicity OLDFIELD * (Secretary)WCMCHuntingdon Road 219CambridgeCB3 ODL UNITED KINGDOM44-1223-277-31444-1223-277-136sara.oldfield@)wcmc.org.uk

MS Peggy OLWELLNational Park ServiceP.O. Box 37127 - MS MIB 3223Washington, D.C.20013-7127 USAl-202-21 9-8933l-202-208-4620Peggy [email protected]

Dr Carlos OSTOLAZAApartado Postal 4338Lima 1PERU

Dr Marisela PONCEFacultad Agronomia UCVVia el LimonMaracay 2101ED0 Aragua4579 VENEZUELA

Mr Walter ROOSLIGsteigstrasse 65Zurich8049 SWITZERLAND41-1-371-737341-1-371-7374

Dr Maurizio SAJEVA +Universita de Palermo, Dip. di ScienzeVia Archirafi 38Palermo90139 ITALY39-91-616-149339-91-617-6089

Dr Uwe SCHIPPMANNScientific Authority to CITESBundesamt fur NaturKonstantinstr. 110Bonn53179 GERMANY49-228-954-343249-228-954-3470

Mrs Meena SINGH +Indian Society of Cacti & SucculentsA-162 Sector 40Noida201303 INDIA

Dr Gideon SMITH *National Botanical InstitutePrivate Bag Xl01Pretoria0001 SOUTH AFRICA27-12-804-320027-12-804-3211

Mr Diedrich SUPTHUT *Staedtische Sukkulentensammlung ZurichMythenquai 88Zurich8002 SWITZERLAND41-l-201-455441-l-201-5540

Dr Nigel TAYLOR * (Chair-elect)Curator of Living CollectionsRoyal Botanic Gardens, KewRichmond, SurreyTW9 3AE UNITED [email protected]

211

Prof Baltasar TRUJILLOBotanical Institute, Agronomy FacultyCentral Univ. of VenezuelaApdo 4579Maracay2101 VENEZUELA58-43-464-56558-43-454-175

Mr Ernst Jacobus VAN JAARSVELDKirstenbosch Botanical GardensPrivate Bag X7Claremont7735 SOUTH AFRICA27-21-762-116627-21-762-3229

Dr Andrew VOVIDESInstituto de Ecologia, A.C.Apdo Postal 63Xalapa, Veracruz91000 MEXICO52-281-8600052-281-86000vovidesaQsun.ieco.conacyt.mx

Dr George WOLSKYSt. Petersburg State UniversityBabushkina Str. 96-55St. Petersburg193012 RUSSIAN FEDERATION7-812-218-96967-812-218-1346telex: 121481 lsu - su

Dr C. R. BABU +Department of BotanyUniversity of DelhiDelhi110007 INDIA

ADDITIONAL ACTION PLAN AUTHORS AND CONTRIBUTORS

Dr David BRAMWELL +Director Jardin Botanic0 ‘Viero y Clavijo’14 Tarifa AltaApdo. De CorreosLas Palmas de Gran Canarias35017 SPAIN

Silvio FICI +Universita de Palermo, Dip. di ScienzeVia Archirafi 38Palermo90139 ITALY

Dr Maike GERBAULET *Bolus HerbariumUniversity of Cape Town7700 SOUTH AFRICA

Steve HAMMER *P.O. Box 72Belen, NM87002 USA

Dr Henk t’HART *Postbus 80.055NL-3508 TB UtrechtNETHERLANDS

Wendy HODGSON *Desert Botanical Garden1201 N. Galvin ParkwayPhoenix, AZ85008 USA

Dr Ulrich MEVE *Botanik Institut und Botanik GartenSchlossgarten 3Munster48149 GERMANY

212

IUCN/SSC Action Plans for the Conservation of Biological Diversity

Action Plan for African Primate Conservation: 1986- 1990. Compiledby J.F. Oates and the IUCN/SSC Primate Specialist Group,1986, 41 pp. (Out of print.)

Action Plan for Asian Primate Conservation: 7987-1997. Compiledby A.A. Eudey and the IUCN/SSC Primate Specialist Group,1987, 65 pp. (Out of print.)

Antelopes. Global Survey and Regional Action Plans. Part 1. Eastand Northeast Africa. Compiled by R. East and the IUCN/SSCAntelope Specialist Group, 1988, 96 pp. (Out of print.)

Dolphins, Porpoises and Whales. An Action Plan for the Conservationof Biological Diversity: 1988- 1992. Second Edition. Compiledby W.F. Perrin and the IUCN/SSC Cetacean Specialist Group,1989, 27 pp. (Out of print).

The Kouprey. An Action P/an for its Conservation Compiled by J.R.MacKinnon, S.N. Stuart and the IUCN/SSC Asian Wild CattleSpecialist Group, 1988, 19 pp. (Out of print.)

Weasels, Civets, Mongooses and their Relatives. An Action Plan forthe Conservation of Mustelids and Viverrids. Compiled by A.Schreiber, R. Wirth, M. Riffel, H. van Rompaey and the IUCN/SSCMustelid and Viverrid Specialist Group, 1989,99 pp. (Out of Print.)

Antelopes. GlobalSurveyand RegionalAction Plans. Part2. Southernand South-central Africa. Compiled by R. East and the IUCN/SSC Antelope Specialist Group, 1989, 96 pp. (Out of print.)

Asian Rhinos. An Action P/an for their Conservation. Compiled byMohd Khan bin Momin Khan and the IUCN/SSC Asian RhinoSpecialist Group, 1989, 23 pp. (Out of print.)

Tortoises and Fresh water Turtles. An Action Plan for theirConservation Compiled by the IUCN/SSC Tortoise andFreshwater Turtle Specialist Group, 1989, 47 pp.

African Elephants and Rhinos. Status Survey and ConservationAction Plan. Compiled by D.H.M. Cumming, R.F. du Toit, S.N.Stuart and the IUCN/SSC African Elephant and Rhino SpecialistGroup, 1990, 73 pp. (Out of print.)

Foxes, Wolves, Jackals, and Dogs. An Action Plan for the Conservationof Canids. Compiled by J.R. Ginsberg, D.W. Macdonald, andthe IUCN/SSC Canid and Wolf Specialist Groups, 1990,116 pp.

The Asian Elephant. An Action P/an for its Conservation. Compiledby C. Santiapillai, P. Jackson, and the IUCN/SSC Asian ElephantSpecialist Group, 1990, 79 pp,

Antelopes. Global Survey and Regional Action Plans. Part 3. Westand Central Africa. Compiled by R. East and the IUCN/SSCAntelope Specialist Group, 1990, 171 pp.

Otters. An Action P/an for their Conservation Compiled by P. Foster-Turley, S. Macdonald, C. Mason and the IUCN/SSC OtterSpecialist Group, 1990, 126 pp.

Rabbits, Hares and Pikas. Status Survey and Conservation ActionPlan. Compiled by J.A. Chapman, J.E.C. Flux, and the IUCN/SSC Lagomorph Specialist Group, 1990, 168 pp.

African lnsectivora and Elephant-Shrews. An Action Plan for theirConservation. Compiled by M.E. Nicoll, G.B. Rathbun and theIUCN/SSC Insectivore, Tree-Shrew and Elephant-ShrewSpecialist Group, 1990, 53 pp.

Swallowtail Butterflies. An Action Plan for their Conservation.Compiled by T.R. New, N.M. Collins and the IUCN/SSCLepidoptera Specialist Group, 1991, 36 pp.

Crocodiles. An Action Plan for their Conservation, Compiled by J.Thorbjarnarson, H. Messel, F.W. King, J.P. Ross and the IUCNISSC Crocodile Specialist Group, 1992, 136 pp.

South American Came/ids. An Action Plan for their Conservation.Compiled by H. Torres and the IUCN/SSC South AmericanCamelid Specialist Group, 1992, 58 pp.

Australasian Marsupials and Monotremes. An Action Plan for theirConservation. Compiled by M. Kennedy and the IUCN/SSCAustralasian Marsupial and Monotreme Specialist Group, 1992,103 pp.

Lemurs of Madagascar. An Action Plan for their Conservation:7993-7999. Compiled by R.A. Mittermeier, W.R. Konstant,M.E. Nicoll, 0. Langrand and the IUCN/SSC Primate SpecialistGroup, 1992, 58 pp. (Out of print.)

Zebras, Asses and Horses. An Action Plan for the Conservation ofWild Equids. Compiled by P. Duncan and the IUCN/SSCEquid Specialist Group, 1992, 36 pp.

Old World Fruit Bats. An Action Plan for their Conservation.Compiled by S. Mickleburgh, A.M. Hutson, P.A. Racey andthe lUCN/SSC Chiroptera Specialist Group, 1992, 252 pp.(Out of print.)

Seals, Fur Seals, Sea Lions, and Walrus. Status Survey andConservation Action Plan. Peter Reijnders, Sophie Brasseur,Jaap van der Toorn, Peter van der Wolf, Ian Boyd, JohnHarwood, David Lavigne, Lloyd Lowry, and the IUCN/SSCSeal Specialist Group, 1993, 88 pp.

Pigs, Peccaries, and Hippos. Status Survey and ConservationAction P/an. Edited by William L.R. Oliver and the IUCN/SSCPigs and Peccaries Specialist Group and the IUCN/SSCHippo Specialist Group, 1993, 202 pp.

The Red Panda, Olingos, Coatis, Raccoons, and their Relatives.Status Survey and Conservation Action Plan for Procyonidsand Ailurids. (In English and Spanish) Compiled by Angela RGlatston and the IUCN/SSC Mustelid, Viverrid, and ProcyonidSpecialist Group, 1994, 103 pp.

Dolphins, Porpoises, and Whales. 7994- 1998 Action Plan for theConservation of Cetaceans. Compiled by Randall R. Reevesand Stephen Leatherwood together with the IUCN/SSCCetacean Specialist Group, 1994, 91 pp.

Megapodes. An Action Plan for their Conservation 1995- 1999.Compiled by Renit W.R.J.Dekker, Philip J.K.McGowan andthe WPA/BirdIife/SSC Megapode Specialist Group, 1995,41 PP*

Partridges, Quails, Francolins, Sno wcocks and Guineafo wl. Statussurvey and Conservation Action P/an 7995-1999. Compiledby Philip J.K. McGowan, Simon D. Dowell, John P. Carrolland Nicholas J.A. Aebischer and the WPA/BirdLife/SSCPartridge, Quail and Francolin Specialist Group, 1995,102 pp:

Pheasants: Status Survey and Conservation Action Plan 1995-7999. Compiled by Philip J.K. McGowan and Pater J. Garsonon behalf of the WPA/BirdLife/SSC Pheasant Specialist Group,1995, 116 pp.

The Wild Cats: Status Survey and Conservation Action Plan.Compiled and edited by Kristin Nowell and Peter Jacksonand the IUCN/SSC Cat Specialist Group, 1996, 406 pp.

Eurasian Insectivores and Tree Shrews: Status Survey andConservation Action P/an. Compiled by David Stone and theIUCN/SSC Insectivore, Tree Shrew and Elephant ShrewSpecialist Group. 1996, 108 pp.

African Primates: Status Survey and Conservation Action Plan(Revised edition). Compiled by John F. Oates and the IUCNISSC Primate Specialist Group, 1996.

Cranes: Status Survey and Conservation Action Plan. Compiledby Curt D. Meine and George W. Archibald and the IUCNISSC Crane Specialist Group, 1’996.

Orchids: Status Survey and Conservation Action Plan. Edited byEric Hagsater and Vinciane Dumont, and compiled by Alec M.Pridgeon on behalf of the IUCN/SSC Orchid Specialist Group,1996, 153 pp.

Palms: their conservation and sustained utilization. Status Surveyand Conservation Action Plan. Edited by Dennis Johnson andthe IUCN/SSC Palm Specialist Group, 1996, 116 pp.

Conservation of Mediterranean Island Plants. I. Strategy forAction. Compiled by 0. Delano& B. de Montmollin, L. Olivierand the IUCN/SSC Mediterranean Islands Plant SpecialistGroup, 1996, 106 pp.

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