Wolfgang Frey (Editor) Syllabus of Plant Families A. Engler , s Syllabus der PÀanzenfamilien 4 Pinopsida (Gymnosperms), Magnoliopsida (Angiosperms) p.p.: Subclass Magnoliidae [Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Asparagales)] B 13 th ed. Borntraeger
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Wolfgang Frey (Editor)
Syllabusof Plant FamiliesA. Engler,s Syllabus der P anzenfamilien
4 Pinopsida (Gymnosperms), Magnoliopsida (Angiosperms) p.p.:Subclass Magnoliidae [Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Asparagales)]
B
13th ed.
Borntraeger
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sPart 4 Pinopsida (Gymnosperms),
Magnoliopsida (Angiosperms) p.p.: Subclass Magnoliidae [Amborellanae to
Half a century ago, the 12th edition (vol. 1, 1954) of Adolf Engler´s well-known “Syllabus der Pflanzenfamilien” (“Syllabus of Plant Families”), ed. by H. Melchior and E. Werdermann was published. Later, a revision of the mosses (13th ed., Kapitel V,2 Bryophytina), by K. Walther, followed in 1983.
The 13th edition will be published in five parts, starting in 2009 with Part 3 “Bryophytes and seedless Vascular Plants” and followed in 2012 by Part 1/1 “Blue-green Algae, Myxo-mycetes and Myxomycete-like organisms, Phytoparasitic protists, Heterotrophic Hetero-kontobionta and Fungi p.p.” and 2015 by Part 2/1 “Eukaryotic Algae [Glaucobionta, Het-erokontobionta p.p. (Cryptophyta, Dinophyta, Haptophyta, Heterokontophyta), Chlorarachniophyta, Euglenophyta, Chlorophyta, Streptophyta p.p.” (except Rhodobi-onta). Now Part 4 “Pinopsida (Gymnosperms), Magnoliopsida (Angiosperms) p.p.: Sub-class Magnoliidae [Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Asparagales)] is ready.
Numerous molecular analyses led to new insights and a better understanding of the evo-lution and systematics of the plant groups and fungi. On the other hand, “classical” morpho-logical and taxonomical expertise is in decline, especially for less showy groups of organisms. As also noted in Part 3, “we are convinced that in the ‘molecular times’ there is an indispen-sable need to preserve the knowledge of the whole diversity and biology of organisms for the next generations. Otherwise, we will not be able to educate experts in the future who will maintain our knowledge of the full range of the earth’s biodiversity”:
The present volume provides a basic treatise of the world-wide morphological and mo-lecular diversity of the Pinopsida (Gymnosperms) and Magnoliopsida (Angiosperms) p.p.: Subclass Magnoliidae [Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Aspara-gales)].
Following the tradition of Engler, and incorporating the latest results from molecular phylogenetics and phylogenomics, the editor and the authors hope to have created an up-to-date overview of families and genera that will serve as reference for a long time.
The authors and the editor are grateful to the publisher, Dr. A. Nägele, for realizing this basic and fundamental systematic treatment, the Syllabus of Plant Families.
The last two and a half decades provided revolutionary new insights into the phylogeny and diversity of organisms on earth. Especially the gymnosperms and the flowering plants have been in the focus of intensive research studies. The relationships were considerably revised.
The most important insights from recent molecular phylogenies and phylogenomic evi-dence allow a new understanding of the relationships between and within the taxa of the gymnosperms and angiosperms and are treated in the view of our decade, including mor-pho-anatomical, molecular and ecological results of the last three decades and bringing them on an up-to date presentation of these plants groups.
The present Part 4 of the 13th edition of “Engler’s Syllabus of Plant Families” gives an up-to-date review of the Pinopsida (Gymnosperms ) and Magnoliopsida (Angiosperms ) p.p.: Subclass Magnoliidae [Amborellanae to Magnolianae , Lilianae p.p. (Acorales to Aspara-gales )] and the relationships on family and generic level, integrating morphological-anatom-ical and molecular data. It provides a thorough treatise of Gymnosperms and of part of the Angiosperms. In the Gymnosperms, the extinct diversity is included, and the present treatise is a first synthesis of classical anatomical-morphological characters with modern molecular data combined with the numerous new discoveries of fossils especially from China made during the last ten years.
The Angiosperms are the most diverse group of plants and form nearly 95% of the global vegetation from arctic tundra, resp. subantarctic vegetation formations, to tropical rainfor-ests. There is actually no comprehensive survey covering all families and genera of angio-sperms. Engler’s Syllabus is an attempt to fill this gap by covering all angiosperms in two volumes arranged according to the most recent phylogenetic system of APG III (2009). In this first volume all families and genera of Magnoliids are described. The monocotyledonous families are covered by the orders Acorales and Alismatales as well as all groups of Liliid orders and families (Petrosaviales , Dioscoreales , Pandanales , Liliales and Asparagales ). Es-pecially Asparagales, comprising c. 40% of all extant monocotyledons, include numerous commercially important ornamental plants in Orchidaceae , Alliaceae and Iridaceae . Also, several economically important species are found in genera like Allium , Aloë, Asparagus , Crocus , and Vanilla . The Orchidaceae are included with a fully revised and modern treatise, thus representing one of the two most species-rich families of plants. The remaining mono-cotyledonous groups (Arecales , Commelinales , Poales , Zingiberales and Dasypogonales ) and the core eudicotyledons will be treated in Part 5.
W. Frey
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(Angiosperms )
Pinopsida (Gymnosperms ) and Magnoliopsida (Angiosperms ) are seed plants producing plants with a sporophyte-dominant alternation of generations. The ancient greek word “gymnospermos” (γυμνόσπερμος) stands for „naked seeds“, αγγείον (angeíon, “receptacle”) and σπέρμα (spérma, “seed”) for angiospermous, seeds within an enclosure. Brongniart (1843) first subdivided the dicotyledons (dicots) in «Angiospermes» and «Gymnospermes». Together with the “Gymnosperms” (Pinopsida) and the Progymnospermopsida the angio-sperms are sister to the ferns (Moniliformopses ) and form the Euphyllophytina .
Extant gymnosperms are trees, shrubs, krummholz (dwarf pine trees) and woody clim-bers; with only one root parasite, Parasitaxus usta (Podocarpaceae ). Especially the tree life form is dominant in the boreal zone of the Northern Hemisphere, e.g., conifers in the boreal forest communities with extremely high economic importance. Due to the small landmasses in the Southern Hemisphere extant gymnosperm-dominated forests cover much smaller ar-eas than those of the boreal forests (“taiga”), but are of great significance for floral history and nowadays considered as relicts of tertiary forest communities (temperate rainforests). The extant gymnosperms are clearly monophyletic and sister to the angiosperms (e.g., Ran et al. 2010), comprising c. 1040 spp. in 4 subclass., 8 ord., and 12 fam. (Christenhusz 2011, Cycadales acc. Osborne et al. 2012). But their relationships to fossil gymnospermous taxa remain obscure and incompletely understood. Especially the position of Gnetidae has been problematical and was controversially discussed [e.g. Qiu et al. 1999: embedded in the coni-fer clade as sister to Pinaceae (“gnepine hypothesis”); Rydin et al. 2002: sisters to all other extant seed plants (cf. Gnetidae)]. The families of modern Pinales appeared in the arid Up-per Permian, major diversification in Early Cretaceous; radiation of most genera within the last 25 mio.y.
The Angiosperms are the most diverse group of plants, comprising all known life forms from trees to annuals and parasites, and form nearly 95% of the global vegetation from arctic tundra, resp. subantarctic vegetation formations to deserts and tropical rainforests. Their evolutionary dominance is mostly due to the double fertilization and their extraordinary vegetative plasticity and the adaptation to numerous and diverse pollinators. Extant angio-sperms comprise c. 250.000–300.000 spp., estimations between 223.000 and 422.125 ; in 2 classes, 18 superord., 68 ord., and 417 fam.
Until 1993 the formal division of angiosperms in dicotyledons and monocotyledons was still undisputed (Kubitzki et al. 1993). An alternative view of relationships among angiosperm groups was that of Huber (e.g., 1991) which ± anticipated the later classifica-
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Table 2-1. Synopsis of classification of the Subkingdom Embryobionta Cronquist, Takht. & W.Zimm. (excl. Streptophyta p.p. , cf. Syllabus Part 2/1).(Cf. also Syllabus of Plant Families, Part 3, 2009; Table 2-1, p. 6.)
Division Marchantiophyta Stotler & Crand.-Stotl. (Hepaticae, Liverworts)Division Bryophyta Schimp. (Musci, Mosses)Division Anthocerotophyta Rothm. ex Stotler & Crand.-Stotl. (Hornworts)Superdivision Polysporangiomorpha Kenrick & P.R.Crane “Protracheophytes ” sensu Kenrick & P.R.Crane (Rhyniophyta H.P.Banks p.p.) Class Horneophytopsida Kenric k & P.R.CraneDivision Tracheophyta Sinnott ex Cavalier-Smith sensu Kenrick &
Gnetidae Pax, Pinidae Cronquist, Takht. & W. Zimm. Class Magnoliopsida Brogn. (Angiosperms ) Subclass Magnoliidae Novák ex Takht. Superorders Amborellanae M.W.Chase & Reveal, Nymphaeanae Thorne ex Reveal, Austrobaileyanae Doweld ex M.W.Chase & Reveal Magnolianae Takht., Lilianae Takht. (Monocots), Ceratophyllanae Takht. ex Reveal & Doweld Subclass Rosidae Takht. (Eudicots) Superorders Ranunculanae Takht. ex Reveal, Proteanae Takht., Trochodendranae Takht. ex Reveal, Buxanae Takht. ex Reveal & Doweld, Myrothamnanae Takht., Dillenianae Takht. ex Doweld, Saxifraganae Reveal, Rosanae Takht., Berberidopsanae Thorne & Reveal, Santalanae Thorne ex Reveal, Caryophyllanae Takht., Asteranae Takht.
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tion based on molecular data (e.g., Chase et al. 1993). Huber proposed a basal division in angiosperms between a “First principal group” comprising magnoliid families, the mono-cotyledons and the Centrospermae , and a “Second principal group” with the rest of the dicotyledons and stated that basal dicotyledonous angiosperms like the magnoliid families are closer related to the monocotyledons than to the rest of the dicotyledons. With mo-lecular data, esp. DNA sequences as a potential source of phylogenetically informative characters, new imput was brought into systematics. Chase et al. (1993) provided a first set of nearly 500 taxa which revealed also a split between Magnoliales , Laurales , the so-called “palaeoherbs ” and the monocots on one side and the modern eudicots with ranunculids, hamamelids, caryophyllids, Rosidae and Asteridae on the other side. Subsequent analyses (e.g., APG III 2009) changed this image slightly but the split between the monosulcate primitive dicots + the monocots and the tricolpate (or with a derived condition) eudicots is still confirmed. (Cf. Chapter 4: “Recent history of angiosperm classification and mo-lecular phylogeny”.)
The sudden appearance and the subsequent dramatic rise of the angiosperms in the Upper Cretaceous and Early Tertiary have caused much debate. The relationships of an-giosperms to the gymnosperms are still a mystery, and it is until now unclear which seed plant fossils belong to the stem-group angiosperms. (Cf. Chapter 4 “Fossil history and evolution”).
Actually no comprehensive survey covering all families and genera of angiosperms is available as the ambitious work of Kubitzki (ref. cited) is still not complete. Engler’s Syllabus intends to be the first comprehensive survey covering all angiosperms in two volumes and arranged according to the most recent phylogenetic system of APG III (2009).
Table 2-1 gives an overview of the classification of the Subkingdom Embryobionta with Pinopsida and Magnoliopsida .
References and further reading
APG III 2009: An update of the Angiosperm Phylogeny Group classification for the orders and fami-lies of flowering plants: APG III. – Bot. J. Linn. Soc. 161: 105–121.
Brongniart, A. 1843: Enumération des genres des plantes cultivés au muséum d’histoire naturelle de Paris suivant l’ordre établi dans l’école de botanque en 1843. – Paris.
Chase, M.W., Soltis, D.E., Olmstead, R.G., Morgan, D., Les, D.H. & al. 1993: Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene rbcL. – Ann. Missouri Bot. Gard. 80: 528–580.
Christenhusz, M.J.M., Reveal, J.L., Farjon, A., Gardner, M.F., Mill, R.R. & al. 2011: A new clas-sification and linear sequence of extant gymnosperms . – Phytotaxa 19: 55–70.
Huber, H. 1991: Angiospermen. Leitfaden durch die Ordnungen und Familien der Bedecktsamer. – G. Fischer, Stuttgart.
Kubitzki, K., Rohwer, J.G. & Bittrich, V. (eds.) 1993: The Families and Genera of Vascular Plants. Vol. II. Flowering Plants. Dicotyledons. Magnoliid, Hamamelidid and Caryophyllid families. Pp. 92–93. – Springer, Berlin.
Osborne, R., Calonje, M.A., Hill, K.D., Stanberg, L. & al. 2012: The worlds list of Cycads. – Mem. New York Bot. Gard. 106: 480–508.
Qiu, Y.-L., Lee, J., Bernasconi-Quadroni, F., Soltis, D.E., Soltis, P.S. & al. 1999: The earliest angio-sperms : evidence from mitochondrial, plastid and nuclear genomes. – Nature 402: 404–407.
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Ran, J.-H., Gao, H. & Wang, X.-Q. 2010: Fast evolution of the retroprocessed mitochondrial rps3 gene in conifer II and further evidence for the phylogeny of gymnosperms . – Mol. Phyl. Evol. 54: 136–149.
Rydin, C., Källersjö, M. & Friis, E.M. 2002: Seed plant relationships and the systematic position of Gnetales based on nuclear and chloroplast DNA: Conflicting data, rooting problems, and the mor-phology of conifers. – Int. J. Plant Sci. 163: 197–214.
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obtuse apex; lamina coriaceous and folded. Permian, Lower Gondwana. – Noeggerathiopsis O.Feistmantel Leaves elongated strap-shaped with sub-parallel venation. Carbonian and Permian, Southern Hemisphere. – Rufloria S.V.Meyen Leaves strap-shaped with parallel venation. One of the most common Angaran foliage types; Pennsylvanian to Permian, Russia. Considered by Anderson et al. (2007) to belong to a separate fam. Rufloriaceae Ledran emend. S.V.Meyen together with Sucho-viella , Pechorostrobus , Krylovia and Cladostrobus . Roots. Amyelon W.C.Williamson ex P.D.W.Barnard Isolated roots which include both protostelic and eustelic forms (“morphospecies”). Prob. produced by cordaite species in response to habitat conditions (Taylor et al. 2009). Endophytic fungi considered as endomycorrhizal are reported from Amyelon-type roots of European Cordaitales (Osborn 1909, Halket 1930). However, it is difficult to decide whether the fungi are true symbionts, parasites or sap-rotrophes (Cridland 1962). Male reproductive organs. Cladostrobus Zalessky, Permian, Russia. – Pe-chorostrobus S.V.Meyen, Permian, Russia. Female reproductive organs. Bardocarpus Zalessky, Per-mian, Russia. – Krylovia Chachlov, Permian, Russia.Fam. Vojnovskyaceae M.F.Neuburg Extinct family based on Vojnovskya M.F.Neuburg. Ovulate stro-bili attached to axis between leafy bracts, ovules reflexed on seed scales. Upper Carboniferous–Lower Permian. Assigned by Taylor et al. (2009) to an order of its own, Vojnovskyales M.F.Neuburg with uncertain affinities.
Paravojnovskya Doweld (3) (Gaussia M.F.Neuburg). Related to Vojnovskya . Reproductive organs a receptacle of c. 2 cm diam., with elongate structures with swollen base and expanded tip on surface. Krassilov & Burago (1981) suggest that this structure is not a strobilus but an uniovulate carpel with style. Early Permian. – Vojnovskya M.F.Neuburg (1). Axis with scars marking the former position of fan-shaped leaves or cataphylls of the Nephropsis -type. Parallel venation. Reproductive organs of fertile branches or cones called “polysperms” (Taylor et al. 2009). Cones of V. paradoxa M.F.Neuburg consist of an axis with attached reflexed seed stalks bearing widened apices: distal portion of cones densely covered by linear scales with interspersed seeds. Seeds of Samaropsis -type. Ovulate cones described as Sergeia neuburgii G.W.Rothwell, G.Mapes & R.H.Mapes, bearing helically arranged scale leaves below and sporophylls above, each bearing a single ovule with integumentary wings. Pollen chamber of ovule simple, with small nucellar beak. Early Permian. – Scirostrobus pterocerum Doweld & S.V.Naugolnykh Stalked, bilaterally symmetrical umbrella-like structure formed by sterile scales and short seed stalks that are fused at base. Vojnovskyalean frutification. Lower Permian, Russia.
Krassilov & Burago (1981) suggested affinities to the angiosperms. Hernandez-Castillo et al. (2003) resolved the Vojnovskyales as basal to Voltziales which form a clade with the early conifers. Mamay (1976) stated that the Vojnovskyales were “a bizarre, short-lived group of late Palaeozoic gymno-sperms”.
2. Order Voltziales Andr. (†)Voltzialean conifers are characterized by orthotropic branches with broad leaves. The dwarf shoots bear flattened and partially fused scales. The “walchian” conifers have plagiotropic branches, needle-like leaves, dwarf shoots with radially arranged scales. Phylogenetic analy-sis (Rothwell et al. 2005) show that the “walchians” are nested within the Voltziales .
Trees, monoecious. Secondary xylem with araucarioid pitting. Leaves needle or scale-like, sometimes furcate at tip. Seed cones consist of an axis with helically arranged bifid bract scales which developed short, uniovulate, radial leafy shoots in their axils. Usually only 1 scale-like leaf was fertile, the remaining being sterile; the fertile appendage or megasporo-phyll was situated near base or middle of the dwarf shoot, bearing a single ovule (cf. Fossil history and evolution of gymnosperms). Stamina composed of a stalk and a flat upper part.
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∞ pollen sacs on the adaxial side of the stalk. Pollen bisaccate (“voltzialean conifers”) or monosaccate (“walchian conifers”). Upper Carboniferous – Lower Jurassic. Phylogenetic analysis (Rothwell et al. 2005) show that the “walchian conifers” are nested within the Volt-ziales .
Historically, Voltziales have been referred as transition between Cordaitales and the modern conifers. The phylogenetic analyses of Hernandez-Castillo et al. (2003) and Rothwell et al. (2005) resolved the Cordaitales and Vojnovskyales as basal to the Voltziales, which form a clade with the early conifers. Within the Voltzialen clade there are 3 additional clades: the Gondwanan Voltziales incl. the Feruglio-cladaceae , the voltzian Voltziales with Majonica , Dolomitia , Aethophyllum , Voltzia hexagona (Bischoff) Geinitz and Ortiseia , and the lebachioid Voltziales with Utrechtia , Otovicia , Emporia, Barthelia and Hanskerpia . The relationships of these taxa with modern conifers are still unclear, and hypotheses in-clude the existence of Cordaitales and modern conifers as separate clades (Doyle 2006) or a polyphy-letic basal grade of conifers (Hilton & Bateman 2006). In earlier treatments, the taxa now assigned to Utrechtiaceae , Majonicaceae and Ullmanniaceae were included in the Lebachiaceae , with the remain-ing morphogenera included in the Voltziaceae (Miller 1977). Giblingodendron nudifolia Falcon-Lang, Kurzawe & Lucas, deciduous coniferopsid, prob. a voltzian conifer with marked tree rings. Macdonal-dodendron giganticus Falcon-Lang, Kurzawe & Lucas interpreted as an evergreen walchian conifer lacking tree rings (Falcon-Lang et al. 2014). Gen. inc. sed.: Shimakuroxylon Philippe, Boura, Oh & al., southernmost Eastern Asia, indicator for warm and wet climates (Philippe et al. 2014).
Fam. Bartheliaceae G.W.Rothwell & G.Mapes Trees? with irregularly branched shoots. Wood with transverse pith septations and sclerotic nests. Leaves up to 5 cm long, either fur-cate or simple (Rothwell & Mapes 2001), with papillate epidermal cells and trichomes at base. Pollen cones small, up to 5 mm long, microsporophylls helically arranged, some with bifid tip. Seed cones compound, occurring in a fertile zone along the axis; each cone radial with numerous sterile scales and ovules orthotropous on narrow sporophylls.
Barthelia G.W.Rothwell & G.Mapes - B. furcata G.W.Rothwell & G.Mapes, Pennsylvanian (Upper Carboniferous), Kansas.Fam. Buriadiaceae D.D.Pant & Nautiyal Branches woody. Leaves helically arranged, poly-morphic, decurrent at base; leaf-tips simple too many times furcate. Stomata absent from abaxial leaf surface. Contrary to the original description (Pant & Nautiyal 1967b) the ovules are not attached along the stem (Singh et al. 2003).
Buriadia Seward & Sahni, Lower Permian, India .Fam. Emporiaceae G.Mapes & G.W.Rothwell Small trees. Ultimate vegetative shoots with simple and/or bifid leaves. Late Pennsylvanian.
Emporia G.Mapes & G.W.Rothwell Ovulate cones with helically arranged bifid bract scales; the axils with a flattened fertile shoot consisting of 20–30 sterile scales and 1–5 fertile scales; each fertile scale with a single inverted ovule attached at apex (Mapes & Rothwell 1990). Seeds with polycotyledonous embryos. - E. lockardii (G.Mapes & G.W.Rothwell) G.Mapes & G.W.Rothwell (Lebachia lockardii G.Mapes & G.W.Rothwell), Permian, Upper Pennsylvanian, Kansas (Hernandez-Castillo et al. 2009) – Hanskerpia G.W.Rothwell, G.Mapes & G.R.Hernandez-Castillo Branches plagiotropic, with simple amphistomatic leaves, c. 10 cm long. Seed cones com-pound, produced on ultimate shoots. Ovules inverted and singly or in pairs on each sporo-phyll. Pollen cones unknown. - H. hamiltonensis G.W.Rothwell, G.Mapes & G.R.Hernandez-Castillo, Late Pennsylvanian, N. America.
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A cladistic analysis (Rothwell et al. 2005) shows Hanskerpia nested within the paraphyletic walchian Voltziales .
Fam. Ferugliocladaceae S.Archangelsky & R.Cunéo Trees with vegetative branches and reproductive organs. Subtending sterile bracts absent. Ovules orthotropous. Early Permian, Gondwana.
Ferugliocladus S.Archangelsky & R.Cunéo Branches with small linear leaves, c. 1 cm long, each with a single vein. Pollen cones terminal; pollen monosaccate. Ovulate cones ter-minal, consisting of helically arranged bract-scales and axillary orthotropous and platysper-mic ovules (Taylor et al. 2009). Seeds with bifid apex. - F. riojanum S.Archangelsky & R.Cunéo, Lower Permian, Argentina. – Genoites Feruglio Branches with helically arranged bifid leaves. Axils of some leaves with stalked orthotropous ovules. Ovulate cones of Ferugliocladus may have originated from reduction of ovulate branches of Genoites. - G. patagonica Feruglio, Per-mian, Argentina. – Ugartecladus S.Archangelsky & R.Cunéo Similar to Ferugliocladus. Seeds lacking a bifid micropyle (Archangelsky & Cunéo 1987). - U. genoensis S.Archangelsky & R.Cunéo, Lower Permian, Argentina.Fam. Majonicaceae Clem.-West. Small plants. Leaves helically arranged, up to 3 cm long, amphistomatic and often heterophyllous. Ovules in lateral to adaxial position on the fertile scale, thus ovules not in a single plane (Clement-Westerhoff 1987, 1988). Acc. to Looy (2007) well differentiated during Early Permian.
Cassinisia H.Kerp & al. Leaves thick, overlapping with falcate tips. Pollen and ovulate cones unknown. - C. orobica Kerp & al. (1996). – Dolomitia Clem.-West. Ovulate cones with c. 13 triangular sterile scales; 3 fertile scales with the 2 lateral ones slightly recurved. Ovules attached to abaxial surface. - D. cittertiae Clem.-West.., Permian, Italy. – Majonica Clem.-West. Short shoots partially fused to the subtending bract-scale and somewhat flattened. - M. alpina Clem.-West., Upper Permian, Italy. – Lebowskia Looy Leaves > 1 cm long, with obtuse apex. A single large median ovuliferous scale flanked by 2 smaller ones (Looy 2007). - L. grandifolia Looy, Late Permian, N America. – Pseudovoltzia Florin Bract-scales and dwarf shoots par-tially fused. Dwarf shoots partially flattened, with 2 sterile and 3 fertile scales; each fertile scale with a single ovule. Ovules not all in the same plane, borne directly on fertile scales and not on stalks. - P. liebeana Florin, Upper Permian, Europe.Fam. Thucydiaceae G.R.Hernandez-Castillo, G.W.Rothwell & G.Mapes Small trees. Wood dense. Pith with resin canals. Stems orthotropic, with plagiotropic lateral branches. Leaves helically arranged on all orders of branching. Ovulate fertile zones between vegetative zones on branch, compound. Pollen cones with helically arranged bracts and axillary dwarf shoots; microsporophylls with 1 terminal erect pollen sac. Ovulate dwarf shoots with sterile scales around base; megasporophylls terminal with recurved apex, each with apical inverted ovule. Late Pennsylvanian.
Thucydia G.R.Hernandez-Castillo, G.W.Rothwell & G.Mapes T. mahoningensis G.R.Her-nandez-Castillo, G.W.Rothwell & G.Mapes, characterized by the combination of an ovulifer-ous fertile zone, compound pollen cones and dissimilar stomatal distribution (Hernandez-Castillo et al. 2001). Late Pennsylvanian (Stephanian), N America.Fam. Ullmanniaceae Nĕmejc Short shoots flattened, with the individual scales fused like ovuliferous scales in extant conifers (Taylor et al. 2009). Vegetative branches with lanceolate-ovate decurrent leaves up to 8 cm long; stomata on both surfaces in irregular rows. Pollen
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cones simple, with abaxially borne elongated pollen sacs. Ovulate cones bearing bract-scales with acute apex. A single ovule on the adaxial surface of the fertile scale.
Ullmannia Göpp. (Cupressites Brogn.). Twigs with cones, Lower Triassic, Germany.Fam. Utrechtiaceae G.Mapes & G.W.Rothwell Forest trees. Stems orthotropic, with plagio-tropic leafy shoots, borne in whorls. Leaves scale-like, helically arranged, only a few mm long, each with a single vein.
Members of this family were formerly included in Lebachiaceae (Florin 1938, 1945) and Walchiaceae (Clement-Westerhof 1984). The name Walchia is still in use for poorly preserved vegetative shoots (Taylor et al. 2009). Utrechtia floriniformis G.W.Rothwell & G.Mapes (Mapes & Rothwell 1991, Roth-well & Mapes 2003) was orig. described to replace the illegitimate name Lebachia .
Ernestiodendron Florin (1). Leaves borne at nearly right angles to the stems. Primary axis of ovulate strobili with helically arranged bifid bract scales. Secondary axes with c. 30 scales, only the distal 4–6 bearing ovules. Cones of E. filiciforme Florin up to 10 cm long, bract scales entire. Lower Permian, Europe. – Moyliostrobus C.N.Miller & J.T.Brown (1). Cones compound, c. 2 cm in diam., with ovate bract scales. Short shoots flattened, with 20–50 sterile bract scales. Each ovule attached to central surface of the cone scale. Integument 3-parted and attached to nucellus only at chalaza. - M. texanum C.N.Miller & J.T.Brown, Lower Permian, N America. – Ortiseia Florin Leaves helically arranged. In O. leonardii Florin, each ovule with 2 horn-like processes at the micropylar end. Pollen described as Nuskiosporites dulhuntyi Potonié, trilete, monosaccate, bell-shaped through distally inclined expansion of sexine. Permian-Triassic, Europe. – Otovicia J.H.F.Kerp & al. Ovulate cones similar to Utrechtia , differing in the shape of the fertile seed scales and the number of ovules per short shoot (Kerp et al. 1990). – Utrechtia G.W.Rothwell & G.Mapes (c. 10) (incl. Lebachia Florin nom. illeg.). Compound ovulate cone of central axis bearing helically arranged bifid bract scales. Axils of each bract scale with short shoot of an axis bearing numerous sterile scales and 1 or more fertile scales. Each fertile scale with single ovule (with micropyle facing the axis) at lateral surface. - U. floriniformis G.W.Rothwell & G.Mapes, Permian, Upper Rotliegend, C Europe.
Ovulate morphogenera. Thuringiostrobus J.H.F.Kerp & Clem.-West. Similar to Walchiostrobus , but differing in the number of fertile and sterile scales. – Walchiostrobus Florin Ovulate cones with a number of sterile scales and 3 fertile scales on each short shoot; each fertile scale with ovule on abaxi-al surface.
Fam. Voltziaceae C.A.Arnold Weakly defined family of Palaeozoic conifers. Taylor et al. (2009) do not accept this taxon but placed their members as Voltzialeans with uncertain af-finities.
Aethophyllum Brongn. Small plants rarely exceeding 2 m. Stems up to 2 cm in diam., with parenchymatous pith and numerous air-spaces (Rothwell et al. 2000). Ovulate cones lax with acuminate bract scales, ovuliferous scales with 5 horn-like projections, each associated with a recurved seed (Grauvogel-Stamm & Grauvogel 1975). Discovery of seedling stage and stem anatomy suggest ecology of a herbaceous ruderal conifer. - A. stipulare Brongn., Lower Triassic Voltzia Sandstone, France. – Conewagia B.J.Axsmith, T.N.Taylor & E.L.Taylor Vegeta-tive shoots with ovulate cones. Leaves helically arranged. Bract-scale complexes flattened, oriented in a single plane and composed of 11 foliar elements. Isolated seeds with poss. wing-
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like structure of the integument. - C. longiloba B.J.Axsmith, T.N.Taylor & E.L.Taylor, Triassic (Up-per Carnian), N America. – Cyadocarpidium Nathorst (c. 5). � cones with helically arranged leaf-like bract scales on short or longer stalks, at base with reduced. Seed scales lobed, bear-ing 2 or 3 inverted, winged or wingless seeds (Farjon 2008). Often associated with leaves of organ gen. Podozamites C.F.W.Braun. Early Triassic to Jurassic, Europe (England, France) to Iran. – Darneya F.Schaarschmidt & Maubeuge Pollen cone, Upper Buntsandstein, France. – Florinostrobus T.Delevoryas & R.C.Hope (Voltzia p.p. ). Seed cones and ovuliferous scales attached to the axis at nearly right angles. Distal portion of ovuliferous scales 5-lobed and bent upwards; adaxial surface with 3 scars prob. from the ovules. - F. andrewsii (T.Delvoryas & R.C.Hope) T.Delevoryas & R.C.Hope, Upper Triassic, N America. Suggested affinties with fam. Ma-jonicaceae (Taylor et al. 2009). – Fredianthus J.M.Anderson & H.M.Anderson (1). Pollen cone, Molteno Formation, S Africa. - F. mayisiformis J.M.Anderson & H.M.Anderson – Glyptolepis W.P.Schimper Morphogenus with bract-scale complexes bearing long cone-scale stalks. Up-per Triassic, C Europe. – Kungurodendron S.V.Meyen Lateral branches plagiotropic to ir-regular. Stomata on upper surface in 2 bands, on lower surface few stomata in groups. Ovu-late cones cylindrical, bract scales simple, with more than 10 sterile scales. Fertile scales c. 10, interspersed with sterile scales. Ovules terminal, inverted. - K. sharovii S.V.Meyen, Permian (Kungurian), Russia. – Lecrosia Florin Lateral branches irregular. Ovulate cones terminal, with simple bract scales, number of sterile and fertile scales unknown. - L. grand’euryi Florin, Carboniferous (Upper Stephanian), France. – Lutanthus J.M.Anderson & H.M.Anderson (3). Pollen cone, Molteno Formation, S Africa. – Sertostrobus L.Grauvogel-Stamm Pollen cone, Upper Buntsandstein, France. – Swedenborgia Nathorst Similar to Tricranolepis , but scale 5-lobed with 5 recurved ovules; the bract scale fused with the fertile scale (Harris 1935). - S. cryptomerioides Nathorst, Rhaetic, Sweden. – Telemachus H.M. Anderson (6) (incl. Odyssian-thus J.M.Anderson & H.M.Anderson, Heidiphyllum Retallack). Seed cone. Molteno Forma-tion, S Africa. -Whole-plant concept and environment reconstruction (Bomfleur et al. 2013). T. elon-gatus J.M.Anderson & H.M.Anderson is thought to represent the seed cone of Heidiphyllum elongatum (Morris) Rettallack (Anderson 1978) and Odyssianthus J.M.Anderson & H.M.Anderson the pollen cone. – Timanostrobus S.V.Meyen Lateral branches irregular. Stomata on upper and lower surface in scattered short rows. Ovulate cones terminal on branches, with more than 10 ster-ile scales; ovuliferous scales interspersed with sterile scales. Ovules terminal, inverted. - T. muravievii S.V.Meyen, Permian (Lower Kazanian), Russia. – Tricranolepis G.Roselt Isolated seed cone with single 3-lobed scale; each lobe fused to the stalk of a recurved ovule (Roselt 1958). - T. monosperma G.Roselt, Triassic (Lower Keuper), Germany. – Voltzia Brongn. Leaves helically arranged and needle-like. Ovulate cones either terminal on branches (V. liebeana Geinitz) or in a fertile zone [V. hexagona (Bischoff) Geinitz]. Ovules inverted on surfaces. Lower Trias-sic. – Voltziopsis Townrow Branches with dimorphic leaves. Ovulate cones at the ends of short branches; each cone consisting of c. 25 bract-scale complexes, ovuliferous scales 5-lobed; each lobe with a single ovule. Permian-Late Triassic, Southern Hemisphere. – Voltz-iostrobus L.Grauvogel-Stamm V. schimperi L.Grauvogel-Stamm, pollen cone, Lower Trias-sic, France – Willsiostrobus L. Grauvogel-Stamm & F. Schaarschmidt (6). Pollen cone, Up-per Buntsandstein, France (Grés à Voltzia).
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Taxa of uncertain affinities to Voltziales Order Dordrechtitales J.M.Anderson & H.M.Anderson (†)Elongate seed cones of numerous subopposite subdecussate fascicles of 3 (–4?). T-shaped ovuliferous scales attached to short pedicels, bract scales absent or fully fused.Fam. Dordrechtitaceae J.M.Anderson & H.M.Anderson (†)
Dordrechtites H.M.Anderson (3). Molteno Formation, Lower Triassic, S Africa. Pollen cones and foliage unknown.
3. Order Pinales Gorozh.Fam. Pinaceae Spreng. ex F.Rudolphi Trees, shrubs or krummholz (Pinus mugo Turra), to 112 m tall (Sequoia sempervirens), evergreen or deciduous (Larix , Pseudolarix ). Monoecious. Highly resinous. Shoots usually with distinctive annual growth and scaly terminal buds. Wood with adaxial parenchyma, with normal (non-traumatic) resin canals and ray trac-
Fig. 3–14. Pinaceae. A–N. Pinus sylvestris. Alternation of generations. A. Branch with pollen cones. B. Pollen cone. C. Microsporophyll, schematic. D–E. Development of microspore (pollen grain) (cf. text), g generative cell. F. Branch with � cones (ovuliferous cone, seed cone). G. � Ovuliferous cone. H. Seed scale with 2 ovules. I. Ovuliferous scale and subtending bract scale, LS. K. Apex of an ovule with pollen tube, integument (i), nucellus (n), embryo sac (e) with prothallium and archegonia (a). L. Seed scale with seed, LS. M. Seed with testa (seed coat) and embryo, LS. N. Germination. R! meiosis, G gametophyte, Sp sporophyte. Without scale. (After Coulter & Chamberlain, Sachs, Strasburger, Wal-ter. Adopted from the 12th ed. of ‘Syllabus der Pflanzenfamilien’ 1954, I, p. 327, fig. 132).
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Fig. 3–15. Pinales. Pinaceae. 1–2. Abies koreana, twig with seed cones. 3. A. alba. Twig with pollen cones. 4. Larix leptolepis. Twig with pollen cones and ovuliferous cones. 5. L. decidua. Twig with pollen cones and seed cones. 6–7. Cedrus atlantica. Twig with seed cones. (Orig.; 1–2, 6–7 W. Barthlott; 3–5 W. Rauh; 1–7 courtesy WB) (Scale bar: 1–2, 6: 5 cm; 3–5: 1 cm; 7: 10 cm).
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heids. Leaves solitary, helically inserted on long or short shoots or arranged in fascicles of 1–8 on dwarf shoots, surrounded by a sheath (Pinus), needle-like to long linear, mostly xero-morphic, each with 1–2 vascular bundles, and 1–many resin canals; palisade parenchyma present or absent. Pollen cones catkin-like, axillary, solitary or clustered, with ∞ helically arranged microsporophylls (each with 2 microsporangia), producing great quantities of pol-len. Pollen [except in Larix (smooth, with a narrow equatorial ridge), Pseudotsuga (smooth, with 3 converging ridges), and Tsuga (with a ring-shaped saccate structure) (non-saccate)] bisaccate or monosaccate (2 or 1 air-bladder). Wind-pollinated. � cones in leaf axils, soli-tary, sessile or shortly stalked, irregulary grouped or in whorls, erect or becoming pendulous. Bract scales and ovuliferous/seed scales helically arranged. Bract scales ± conspicuous in ovuliferous cones, often reduced in mature, woody seed cones. Ovuliferous/seed scales in the axil of bract scales, never fused with bract scales, on the adaxial (upper) side of each fer-tile scale bearing 2 inverted ovules/seeds. Seed cones erect or pendulous, woody, small to moderate or occ. of great size, matering in 1–2(–)3 years. Mature cones with woody axis (breaking up in Pseudolarix ). Bract and seed scales both persistent or deciduous in Abies and Cedrus . Development of wings by the seed scale. Seeds slightly flattened, with 1 wing or wingless (few species of Pinus). Seeds wind-dispersed; in few species (cf. Pinus ) dispersal by birds (ornithochory, dysochory). 2n = 24 (in Pseudolarix 2n = 44, Pseudotsuga menziesii 2n = 26). Exclusively N hemispheric family (Laurasian), except P. merkusii Jungh. & de Vriese, northern Sumatra and in the Philippines, crossing the equator. Predom. in boreal Eurasia and N America; absent from trop. lowlands; in arid regions only some Pinus spp. 4 major centres of diversity: Sino-Himalayan region, Japan with Taiwan, California and Mexico. Spe-cies of Abies, Larix, Picea and Pinus are reaching the timberline in the high mountains and the northern limit of trees to the Arctic tundra. Adaptation to fire in Pinus. P. longaeva (Inner mt. ranges, Utah, Nevada, E California, USA), c. 4800 y. Of great economic importance. No fossils known from S Hemiphere. 11 gen. (231). Figs 3-14–3-17.
Subdivided in 3 subfam., Pinoideae Pilg. (Pinus , Cathaya , Picea ), Laricoideae Melch. & Werdermann (Larix , Pseudotsuga ) and Abietoideae Pilg. (Abies , Cedrus , Keteleeria , Nothotsuga , Pseudolarix , Tsuga ) (cf. Farjon 2010). Cf. molecular phylogenetic analysis by Liston et al. (2003). “The gen. Cedrus has come out as basal (sister group) to all other genera of the Pinaceae in some recent DNA-based cladistic analyses; morphological evidence however places it firmly within Abietoideae. A basal position is not corroborated by the fossil record. Its position is tentative” (Farjon 2010, p. 38). Comparative wood anatomy in Abietoideae (Esteban & de Palacios 2009).Fossil history. Origin in the arid Upper Permian, major diversification in Early Cretaceous; radiation of most gen. within the last 25 mio.y. (Leslie et al. 2012, Farjon & Filer 2013). Ductoabietoxylon Kur-zawe & Merlotti, Scleroabietoxylon Kurzawe & Merlotti (both genera with abietoid pitting), Parnai-boxylon Kurzawa & Merlotti and Damudoxylon Maheshwari, Kaokoxylon Kräusel and Taeniopitys Kräusel (the latter 3 with araucarian pitting) from the Permian permineralized flora of the Parnaíba
Fig. 3–16. Pinales. Pinaceae. 1–2. Pseudotsuga menziesii. Seed cones with bract scales and seed scales. 3. P. canadensis. Twig with ovuliferous cones. 4–5. Picea abies. 4. Twig with seed cones. 5. Habit. 6. P. orientalis. Twig with pollen cones. 7. Abies homolepis (left), Picea abies (right). Twig with needles; needles with cushions. (Orig.; 1–2 phot. W. Rauh; 3, 7 W. Barthlott; 4–6 E. Fischer; 1–3, 7 courtesy WB) (Scale bar 1, 3, 6–7: 1 cm; 2: 5 mm; 4: 5 cm; 5: 5 m).
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Phylogeny. Posidoniaceae is part of the Cymodoceaceae -compl ex of Les et al. (1997) and is closely related to Cymodoceaceae and Ruppiaceae .
Fam. Potamogetonaceae Bercht. & J.Presl. nom. cons. (incl. Hydrogetonaceae Link, Zan-nichelliaceae Chev. nom. cons.) Perennial or annual floating-leaved or submerged freshwa-ter to brackish water herbs. Lower stems rhizomatous or stoloniferous. Leaves alternate, heli-cal, the upper leaves subopposite, opposite throughout or pseudoverticillate, sessile to petiolate, entire to serrulate; ligules forming a tubular sheath; lamina in submerged leaves thin, linear to orbicular, in floating leaves often coriaceous, lanceolate to elliptic or ovate. Inflorescence pedunculate, axially or terminal, with 2 opposite flowers or with more than 2 flowers in a capitates or interrupted spike with a subtending spathe or inflorescence prolifer-ating sympodially (Zannichellia -group ). Flowers monoecious to dioecious, staminate flow-ers with perianth absent or with 3 tepals. Stamens 1. Anthers 2–8-sporangiate. Carpellate flowers with tubular perianth of 3–4 segments, with 1–8 carpels; stigma enlarged, peltate (Zannichellia-group), or flowers hermaphroditic, hypogynous, polysymmetric; tepals (2)–4, greenish, reddish or brownish; stamens 4, rarely 2–3; filaments adnate to perianth; gynoe-cium of (1–)4(–8) carpels, ± stipitate, partly ascidiate, 1-locular; ovules single (Potamogeton -group ). Fruits usually drupelets with membranous exocarp, fleshy mesocarp and stony en-docarp, rarely berries. Seeds without endosperm at maturity. x = 7, 12, 14–18. Cosmopol., diversity centers in N temp. hemisphere. 7 gen. (126). Fig. 4-30.2–5.Potamogeton -group . Groenlandia J.Gay (1). Perennial freshwater herbs. Leaves submerged, opposite, sessile, linear-lanceolate to broadly lanceolate. Inflorescence submerged or held slightly above water surface. - G. densa (L.) Fourr., Spain to Norway, Algeria, Siberia and Afghanis-tan. – Potamogeton L. (100). Annual or perennial freshwater herbs. Leaves alternate to sub-opposite, floating leaves coriaceous, submerged leaves membranous; stipules of submersed leaves, if fused adnate to lamina less than ½. Inflorescence peduncle stiff, not flexible, pro-jecting above water surface. Cosmopol. – Stuckenia Börner [incl. Coleogeton (Rchb.) D.H.Les & R.R.Haynes] (11). Mainly annual freshwater herbs. Leaves submerged, alternate, sessile, linear; stipules tubular, adnate to lamina for at least 2/3. Inflorescence a capitate or cylindri-cal spike, peduncle flexible. Cosmopol.Zannichellia -group . Althenia F.Petit (2). Annual or perennial herbs of coastal brackish water. Leaves distichous, stipulate, narrow, less than 1 mm wide. Inflorescence sympodial. Perianth of 3 minute scales or teeth. Fruit a drupelet. S Europe, N Africa. – Lepilaena Harv. (5). Annual or perennial freshwater plants. Leaves distichous, less than 1 mm wide. Inflores-cence sympodial. Perianth scales minute. Fruit a nutlet. Australia, New Zealand. – Pseud-althenia (Graebn.) Nakai (1). Annual herb in seasonal watercourses. Leaves distichous, usu-ally more than 1 mm wide. Inflorescence sympodial. Fruit a drupelet. - P. aschersoniana (Graebner) Hartog, S Africa. – Zannichellia L. (6). Annual or perennial freshwater or brackish water herbs. Leaves pseudoverticillate, in whorls of 3, less than 1 mm wide. Inflorescence with 1 staminate and 1 carpellate flower. Nearly cosmopol. - Detailed descriptions: Haynes et al. (1998e, h).
Phylogeny. Potamogetonaceae are sister to Zosteraceae (s. bl.), and the clade formed by these 2 fami-lies is strongly supported in the analysis of Les et al. (1997). The morphologically very distinctive Zan-nichellia is rather weakly embedded within Potamogetonaceae (Les et al. 1997) but is here included following APG III (2009) and Les & Tippery (2013). The genera Althenia , Lepilaena and Pseudalthe-
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nia , traditionally placed in Zannichelliaceae have yet not been available for a molecular study (Les & Tippery 2013). Potamogeton is resolved as monophyletic if Stuckenia (incl. Coleogeton , Les & Haynes 1996) is excluded. For a revision of Stuckenia see Kaplan (2008).
Fam. Ruppiaceae Horan. nom. cons. Annual or rarely perennial submerged freshwater or brackish water herbs. Leaves alternate to subopposite, divided into lamina and stipular au-riculate sheath; lamina entire below, serrulate above, linear. Inflorescence a 1-few-flowered capitate axillary or terminal spike, first subtended by 2 subopposite leaves, often with elon-gating peduncle. Flowers perfect, hypogynous, polysymmetric. Perianth absent. Stamens 2, sessile; thecae 2-sporangiate. Carpels (2–)4(–16), each with sessile, peltate stigma. Ovules 1 per carpel, pendulous, campylotropous. Fruits long-stipitate or sessile drupes. Seeds without endosperm. x = 8–12, 15; dimorphic. 1 gen. (4).
Ruppia L. (4). Characters of the family. Cosmopol. - Detailed descriptions: Haynes et al. (1998f) and Ito et al. (2010).
Phylogeny. Ito et al. (2010) looked at the biosystematics of this difficult group in which cytological variation is considerable and both hybridisation and long distance dispersal are likely. They resolved 3 distinct species and a species complex (R. maritima L. s.l.) which possibly contains up to 6 lineages. Ruppiaceae are closely related to Posidoniaceae and Cymodoceaceae (Les et al. 1997), and are even included in the latter by Les & Tippery (2013). We follow APG III (2009) in maintaining the mo no-typic Ruppiaceae.
Fam. Scheuchzeriaceae F.Rudolphi, nom. cons. Perennial herbs with sympodial rhizome, covered with fibrous remains of old leaves. Leaves alternate, distichous, linear, compressed-terete, with sheating base; apex obtuse with a distinct pore; sheath with 2 obtuse auricles. Inflorescence a bracteate raceme with a terminal flower. Flowers perfect, hypogynous, trime-rous and pentacyclic. Tepals 6, lanceolate-elliptic. Stamens 6, anthers bithecate, 4-sporangi-ate. Carpels 3(–6), only basally connate, with decurrent sessile stigmas. Ovules (1–)2 per carpel, erect, anatropous. Fruits follicles. Seeds without endosperm. x = 11. 1 gen. (1). Fig. 4-31.1.
Scheuchzeria L. (1). Characters of the family. - S. palustris L., Sphagnum bogs in temp. to subarctic regions of the Holarctic, S to N California, New Jersey, Italy, Croatia, Japan. - Detailed de-scription: Haynes et al. (1998g).
Phytochemistry. Scheuchzeria is rich in the cyanogenic glucoside triglochinin. Flavonoids, saponins and alkaloids are absent (Haynes et al. 1998g).
Fam. Tofieldiaceae Takht. Perennial herbs with rhizome. Leaves basal, usually distichous, equitant and unifacial. Inflorescence a raceme or rarely a spike, uniflorous in Harperocalis flava McDaniel. Flowers usually hypogynous, polysymmetric. Bracts present. Bracteoles of-ten calycular, forming the calyculus below individual flowers. Tepals 6, in 2 whorls, free, petaloid, usually white to greenish-yellow. Stamens usually 6 in 2 whorls, rarely 9 or 10 (Pleea ). Ovary 3-carpellate; carpels connate only at base, stipitate. Septal nectaries or tepal
Fig. 4-30. Alismatales. Posidoniaceae. 1. Posidonia oceanica. Habit. Potamogetonaceae. 2. Potamo-geton natans. Habit. 3. P. polygonifolius. Habit. 4–5. P. schweinfurthii. Inflorescence. (Orig.; 1 phot. W. Barthlott, 2–5 phot. E. Fischer; 1 courtesy WB) (Scale bar 1–5: 1 cm).
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nectaries usually present. Ovules anatropous or campylotropous, numerous per locule, pla-centation parietal. Fruit a septicidal capsule. x = (14)15 (16). Temp. Eurasia, N to S America. 4 gen. (31). Fig. 4-31.2–4.
Harperocallis McDaniel (incl. Isidrogalvia Ruiz & Pav.) (14). Perennial herbs. Leaves mostly basal, distichous. Inflorescence a raceme or flower solitary, terminal (H. flava McDan-iel). Flowers cup-shaped, white to cream or greenish. Calycular bracteoles 3, free. Seeds with a white chalazar appendage. W Florida, S America (Venezuela to Colombia, Ecuador, Bo-livia). As Harperocallis has priority over Isidrogalvia, Campell & Dorr (2013) transferred all species of Isidrogalvia to Harperocallis. – Pleea Michx. (1). Perennial herb. Leaves basal, distichous, equi-tant, linear. Bracts spathe-like, bracteole 3-partite. Stamens 9, outer whorl 6, inner whorl 3. - P. tenuifolia Michx., coastal plains, SE USA (N Carolina to Florida). – Tofieldia Huds. (12). Peren-nial herbs. Stems glabrous. Leaves basal, equitant. Inflorescence racemose, open or dense, spike-like. Flowers greenish to white. Calycular bracteole united, often present. Seeds with either short chalzal and micropylar appendages, or appendages lacking. Temp. to subarctic regions of the Northern Hemisphere. – Triantha (Nutt.) Small (4). Perennial herbs. Stems glandular-pubescent. Leaves basal, equitant. Inflorescence racemose, open or dense, spike-like. Flowers white. Seeds with long filiform chalzal appendages, micropylar appendages sometimes lacking. N America (3), Japan [1, T. japonica (Miq.) Baker].
Morphology. Comparisons between the calyculus of Tofieldiaceae , formed by 2 or 3 connate scales, with the spathe of Hydrocharitaceae and pseudowhorls of bracts in Alismataceae have been made (Remizowa & Sokoloff 2003, Remizowa et al. 2006a). Remizowa et al. (2010a) clarify the vasculature of this structure, which is usually supplied by 3 vascular traces, but in Tofieldia pusilla (Michx.) Pers. it is basal on the pedicel and supplied by a single trace. The nectaries of Tofieldiaceae are unique to the group; they are triradiate, borne on the inner bases of the connate carpellary stipes (Remizowa et al. 2006b).Phytochemistry. Steroidal saponins and chelidonic acid are present.Phylogeny and classification. Tofieldiaceae were only recently described by Takhtajan in 1995 (Takhtajan 2009), and are still included in Nartheciaceae by Tamura (1998d). Earlier classifications (e.g., Hutchinson 1934) placed the genera even at the base of a broadly defined Liliaceae . Molecular analyses showed that Nartheciaceae sensu Tamura (1998d) are polyphyletic (Chase et al. 1995), and that Tofieldia and Pleea are sister to Alismatanae and Aletris and Narthecium close to Dioscoreaceae (s. bl.). Azuma & Tobe (2011) showed that Pleea is sister to rest of the family, in which Tofieldia is sister to Triantha . Harperocallis , considered as sister to Isidrogalvia by Azuma & Tobe (2011) was shown to be congeneric (Remizowa et al. 2011). For descriptions of Isidrogalvia see Campbell 2010, Cruden 1991 and Cruden & Dorr 1992. Azuma & Tobe (2011) also confirmed that Tofieldiaceae are a well-supporte d monophyletic clade in the order Alismatales .
Fam. Zosteraceae Dumort. nom. cons. Dioecious (Phyllospadix ) or monoecious (Zostera ). Perennial or rarely annual marine submerged herbs with monopodial or rarely sympodial rhizomes. Leaves alternate, distichous, with distinct lamina and sheating base; leaf sheath ligulate, auriculate; lamina linear stomata absent. Inflorescence on erect annual shoots.
Fig. 4-31. Alismatales. Scheuchzeriaceae. 1. Scheuchzeria palustris. Habit. Tofieldiaceae. 2–4. Tofiel-dia calyculata. 2. Habit. 3. Inflorescence. 4. Detail of inflorescence. Zosteraceae. 5. Phyllospadix spec. Habit. (Orig.; 1–5 phot. E. Fischer) (Scale bar 1–3: 1 cm, 4: 1 mm, 5: 5 cm).
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Flowers arranged in 2 rows on one side of a flattened spike (spadix), surrounded by a modi-fied leaf sheath (spathe). Perianth lacking. Staminate flowers with retinacules (possibly modi-fied perianth), 1 stamen, with 2 free 2-locular thecae. Carpellate flowers with 1 unilocular carpel bearing 2 long stigmas. Ovule 1 per carpel, pendulous, atropous. Fruit a nut. x = 6, 9, 10. Temp. to subtrop. marine habitats. 2 gen. (22). Fig. 4-31.5.
Phyllospadix Hook. (6). Dioecious. Rhizome monopodial, thick. Leaves 1 per node, ligu-late, auriculate. Inflorescence with stalked spathe resembling vegetative leaves. Staminate spathe with retinacules inserted alongside flowers. Carpellate spathes with flowers alternat-ing with retinacules. On rocks, temp. northern Pacific Ocean. – Zostera L. (incl. Heterozos-tera Hartog) (16). Monoecious. Rhizome monopodial or rarely sympodial, slender. Leaves 2–6 per shoot, ligulate, auriculate. Spathe stalked, with alternating staminate and carpellate flowers. On muddy or sandy substrates, temp. Eurasia, E Africa, Australasia, Tasmania, N Chile. - Detailed description: Kuo & McComb (1998c).
Pollination. Intertidal species of Zostera and Phyllospadix are epihydrophilous, and pollen is released on the water surface where it forms a network (Cox 1988). In subtidal species the pollination must be hypohydrophilous, and the filiform pollen is wrapped around the stigma (Kuo & McComb 1998c).Phylogeny. Zosteraceae are sister to Potamogetonaceae (Les et al. 1997, Les & Tippery 2013). Only 2 sister genera (Phyllospadix , Zostera ) are recognized as Heterozostera was resolved as being nested with-in Zostera (Les et al. 2002b). The genus Zostera has been studied by Kato et al. (2003).
3. Order Petrosaviales Takht. (incl. Miyoshiales Nakai, Petrosaviineae Shipunov)1 fam., 2 gen. (4).Fam. Petrosaviaceae Hutch. nom. cons. (incl. Japonoliriaceae Takhtajan, Miyoshiaceae Na-kai) Perennial herbs. Achlorophylous and mycoheterotropic (Petrosavia ) or autotrophic (Ja-ponolirion ). Leaves absent, reduced to alternate scales (Petrosavia) or basal, linear, helical, bifacial (Japonolirion). Inflorescence racemose, bracteate. Flowers polysymmetric, with 6 tepals, outer more narrow than inner tepals. Stamens 6, filaments partly adnate to carpels and inner tepals (Petrosavia) or free (Japonolirion). Carpels 3, connate at base, superior, semi-inferior in Petrosavia sakuraii, or free but loosely connected by interdigitating papillae; styles 3, with subcapitate (Petrosavia) or decurrent (Japonolirion) stigma. Ovules numerous (Petrosavia) or 4 per carpel (Japonolirion). Fruit a septicidal capsule or with follicles. x = 12, 13, 15. SE Asia, China, Japan. 2 gen. (4).
Japonolirion Nakai (1). - J. osense Nakai, Japan, on serpentine in Hokkaido and Honshu. – Pe-trosavia Becc. (incl. Protolirion Groom, Miyoshia Nakai) (3). Myanmar, Thailand, Vietnam, Malay Peninsula, N Sumatra, the Philippines, E China, Taiwan, Japan. - Detailed description: Cameron et al. (2003).
Morphology and anatomy. Remizowa et al. (2010b) and Remizowa (2011) suggest that the position of the septal nectaries in both the ascidiate and plicate zone of the gynoecium might be unique and thus a synapomorphy for the Petrosaviales-clade. The embryology of Japonolirion and Petrosavia has been studied by Tobe (2008) and Tobe & Takahashi (2009) who showed that the anatropous and crassi-nucellate ovule with glandular tapetum, a 2-cell-layered nucellar cap formed early in development, the cellular mode of endosperm formation and exotegmic seeds clearly differs from basal (Acoraceae , Araceae ) and more derived monocotyledons (Nartheciaceae , Velloziaceae , Triuridaceae ). Thus a dis-tinct position for Petrosaviales is supported.
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Systematics and phylogeny. Cameron (2007) provides a summary of phylogenetic studies on the family. Dressler (1993) recognized 5 subfamilies of Orchidaceae mainly based on floral morphology: Apostasioideae , Cypripedioideae , Spiranthoideae , Orchidoideae , and Epidendroideae (incl. tribe Va-nilleae ). First phylogenetic analyses based on molecular data (Cameron et al. 1999) mainly confirmed these subfamilies. However, tribe Vanilleae was raised to subfamiliar rank, and subfam. Spiranthoi-deae was reduced to a tribe of Orchidoideae. Molecular data also confirmed the broad circumscription of Orchidaceae incl. the former families Apostasiaceae and Cypripediaceae . This subdivision of Or-chidaceae was also confirmed by Freudenstein et al. (2004). Numerous modern analyses mainly based on molecular data are available, which sometimes considerably altered the traditional circum-scription of taxa. An updated classification was provided by Chase et al. (2015). The following ex-amples of groups that have been dealt with are listed here: Apostasioideae (Kocyan et al. 2004), Cypripedium (Li et al. 2011), Orchidinae (incl. Habenariinae ) (Bateman et al. 2003, Jin et al. 2014), Habenaria (Batista et al. 2013), Orchidinae (Bateman et al. 2009), Ophrys (Soliva et al. 2001), Diseae (Douzery et al. 1999), Disa (Bytebier et al. 2007, 2008), Pogonieae (Pansarin et al. 2008), Cranich-ideae , Spiranthinae (Salazar et al. 2003, Salazar et al. 2009), Discyphinae (Salazar et al. 2014), Cala-deniinae and Drakaeinae (Hopper 2009), Arethuseae (Goldman et al. 2001), Cymbidium (Sharma et al. 2012), Maxillarieae (Whitten et al. 2000), Laeliinae (van den Berg et al. 2009), Oncidiinae (Neubig et al. 2012), Telipogon (Williams et al. 2005), Vandeae (Carlsward et al. 2006), Aeridinae (Zhang et al. 2013). Carlsward et al. (2006) showed that subtribes Aerangidinae and Angraecinae are polyphyletic individually, but together they form a well-supported monophyletic group in all molecular analyses. Therefore, they proposed recognizing a broadly circumscribed subtribe Angraecinae Summerh. that includes Aerangidinae Summerh. Gardiner et al. (2013) provided a phylogeny of Vanda and pro-posed to include Ascocentrum , Euanthe , Christensonia , Neofinetia , Trudelia , and Aerides flabellata Rolfe ex Downie into this genus.Economic importance. Vanilla planifolia Andrews is the only orchid of significant economic impor-tance as an edible crop (Fouché & Jouve 1999). Major vanilla cultivating countries are (amount in tons per year acc. to Fouché & Jouve 1999): Madagascar (700 t), Indonesia (400 t), Comoro Is. (150 t), Ré-union (12 t), Mauritius, Tonga, Tahiti (here Vanilla tahitiensis J.W.Moore), Mexico, China (each less than 50 t per year). Vanilla is the second most expensive spice in the world after saffron (Crocus sativus L., Iridaceae , s. ab.). However, the economic importance of orchids as indoor plants can not be under-estimated. In 2009, c. 45.600.000 potted orchids have been sold only in Germany, and the customers paid c. 400.000.000 € (AMI Agrarmarkt Informationsdienst http://ami-informiert.de). Among these orchids are mainly hybrids of Phalaenopsis , followed by Cymbidium , Dendrobium , Miltonia , Oncidium and Paphiopedilum .
Fam. Ruscaceae M.Roem. nom. cons. (incl. Aspidistraceae Endl., Convallariaceae Horan., Dracaenaceae Salisb., Eriospermaceae Endl., Nolinaceae Nakai, Ophiopogonaceae Endl., Peliosanthaceae Salisb., Polygonataceae Salisb., Sansevieriaceae Nakai, Tupistraceae Schniz-lein) Herbs, shrubs, trees or climbers. Leaves well-developed or scarious, helical, distichous or rarely opposite [Polygonatum oppositifolium (Wall.) Royle]. x = 5–7, 9, 18–21. Circumbo-real to trop., diversity centers in trop. Africa, C Asia, SE Asia, Western N America. For de-scriptions see subfamilies. 25 gen. (730). Figs 4-100–4-102.1.Subfam. Convallarioideae Herb. (incl. Aspidistraceae Endl., Convallariaceae Horan., Ophi-opogonaceae Endl., Peliosanthaceae Salisb., Polygonataceae Salisb., Tupistraceae Schnizlein) Terrestrial or rarely epiphytic herbs with subterranean or exposed rhizomes. Leaves alter-nate, distichous, opposite or verticillate, sessile or petiolate. Inflorescence a terminal or axil-lary raceme or spike, or terminal reduced dibotrya, or flowers solitary. Flowers radially sym-metric to slightly monosymmetric. Tepals 6, in 2 whorls, rarely 4–5 or 2 dimerous whorls,
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sometimes connate to a tube. Stamens 6, rarely 4, 8, 10 or 12 in 2 whorls, filaments free or connate. Ovary superior, 2–5-locular. Ovules anatropous, campylotropous or atropous. Fruit a berry, dry dehiscent capsule (Rohdea ), drupe (Tupistra ) or dry irregularly rupturing cap-sule (Liriope , Ophiopogon , Peliosanthes ).
Tribe Convallarieae Dumort. Monopodial rhizomes. Inflorescence axillary, racemose or spicate. Pedicels articulated (Convallaria , Speirantha , Theropogon ) or not. Tepals connate (Convallaria) or free. Anthers adnate to tepals (Convallaria) or free. Ovary 3-locular, rarely 3–4-locular (Rohdea p.p., Tupistra p.p.) or 1-locular (Tupistra p.p.). Fruit a berry.
Aspidistra Ker Gawl. (98). Acaulescent herbs. Leaves solitary or in fascicles, or 2–4, ovate-elliptic to linear-lanceolate, petiolate. Flowers axillary, solitary, campanulate to urceolate. Tepals 6–8, rarely 4–10, fleshy, connate, purple, pale yellow or white with purple spots. Sta-mens (4–)6–12, attached a base of tepals, filaments short or stamens sessile. Carpels 3–5. Ovary 3–5-locular. Ovules 1 to numerous per locule. Fruit a blue, black or green berry. Hi-malayas, Indochina, S China to SW Japan. – Convallaria L. (3). Acaulescent herbs. Leaves 2, distichous, ovate-elliptic, forming an aerial pseudostem. Inflorescence racemose. Flowers campanulate, nodding. Tepals 6, connate, white. Ovules 4–8 per locule. Fruit a red berry. Circumboreal, sometimes considered as 1 variable sp. – Reineckea Kunth (1). Acaulescent herbs. Leaves distichous, sessile, linear to lanceolate. Inflorescence spicate. Tepals 6, connate, lobes reflexed, pale purplish red. Stamens 6. Ovary 3-locular. Ovules 2 per locule. Fruit a red berry. - R. carnea (Andrews) Kunth, China, Japan. – Rohdea Roth (incl. Campylandra Baker, Gonioscypha Baker) (17). Acaulescent herbs. Leaves ± rosulate, sessile to petiolate, ovate-lanceolate. Inflorescence spicate. Tepals 6, fleshy, connate, yellow, white, green to orange. Stamens 6, attached to tepals. Ovary 3–4-locular. Ovules 2 – 4 per locule. Himalayas, Indo-china, China, Japan. – Speirantha Baker (1). Acaulescent herbs. Inflorescence racemose. Leaves rosulate, ovate-lanceolate, petiolate. Tepals 6, free, white. Ovules 3–4 per locule. - S. gardenia (Hook.) Baker, SE China. – Theropogon Maxim. (1). Acaulescent herbs. Inflorescence racemose. Leaves distichous, sessile, linear. Flowers campanulate, nodding. Tepals 6, free, outer tepals nectariferous. Ovules 6–10 per locule. Fruit a black berry. - T. pallidus (Wall. ex Kunth) Maxim., Himalayas to SW China. – Tupistra Ker Gawl. (incl. Tricalistra Ridl.) (20). Acaulescent herbs. Leaves alternate, ovate to lanceolate, petiolate. Inflorescence spicate. Te-pals 6 or 8, connate, dull yellow to purplish. Stamens 6 or 8. Carpels 3–4. Ovary 1–4-locular. Ovules 2–4 per locule. Fruit a berry or drupe. E Himalayas, Malesia, S China. — Tribe Ophi-opogoneae Endl. Rhizomes spreading, sympodial. Inflorescence terminal, reduced to dibot-rya or racemes. Pedicels articulated. Flowers hypogynous (Liriope ) or perigynous (Ophiopo-gon , Peliosanthes ). Ovary 3-locular. Fruits dry, rupturing irregularly. Seeds fleshy. – Liriope Lour. (6). Acaulescent caespitose herbs. Leaves alternate, sessile, linear to lanceolate. Inflores-cence a reduced dibotryum. Flowers slightly monosymmetric. Tepals 6, free or basally con-nate, white or violet. Stamens 6, attached basally to tepals. Ovules 2 per locule. Seeds black. Indochina, Philippines, China, Japan. – Ophiopogon Ker Gawl. (67). Acaulescent or caules-cent caespitose herbs. Inflorescence a reduced dibotryum or raceme. Tepals 6, basally con-
Fig. 4-100. Asparagales. Ruscaceae. 1–2. Convallaria majalis. 1. Habit. 2. Detail of inflorescence. 3. Polygonatum odoratum. Flowers. 4. Ruscus colchicus. Phylloclades with flower and fruit. (Orig.; 1–4 phot. E. Fischer) (Scale bar 1, 3–4: 1 cm, 2: 5 mm).
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nate or free, white to violet. Ovules 2–6 per locule. Seeds fleshy, blue. India, the Himalayas, Indochina, Malesia, China to Japan. – Peliosanthes Andrews (18). Acaulescent or caulescent caespitose herbs. Inflorescence a reduced dibotryum or raceme. Tepals 6, connate for 1/3–2/3. Stamens 6, filaments short, attached to an expanded, fleshy, epitepalous corona. Ovules 1–5 per locule. Seeds fleshy, blue. India, Indochina, Malesia, S China, Taiwan. — Tribe Po-lygonateae Benth. Rhizomes sympodial. Inflorescence a terminal simple or compound ra-ceme, or condensed racemes or flowers singly in leaf axils. Tepals free or fused. Anthers free or adnate to the tepals. Ovary 3-locular, rarely 2-locular (Maianthemum p.p.). Fruit a berry. Seeds usually pale brown. – Disporopsis Hance (7). Caulescent herbs. Leaves lateral and pseudoterminal, distichous. Flowers axillary, fasciculate or solitary. Flowers campanulate to funnel-shaped, erect or nodding. Tepals 6, connate for ½ of their length, white or yellowish. Stamens 6, connate to tepals by a corona at apex of perigone tube. Ovules 4–6 per locule. Fruit a dark bluish purple berry. Indochina, Philippines, S China, Taiwan. – Heteropolygo-natum M.N.Tamura & Ogisu (6). Caulescent epiphytic herbs. Leaves lateral, distichous, ± petiolate. Inflorescence axillary or terminal, 1-, 2- or 3–6-flowered. Flowers tubular to cam-panulate, nodding. Tepals 6, connate for ½ of their length, corona absent. Stamens 6, outer 3 stamens shorter than inner. Fruit an orange berry. China. – Maianthemum Wigg. (incl. Smi-lacina Desf.) (39). Caulescent herbs. Leaves lateral, distichous, ± petiolate, ovate to lanceolate or sagittate. Flowers usually trimerous, sometimes dimerous, bi- to unisexual. Tepals 4 or 6, free or connate, white, yellow or purplish brown. Stamens 4 or 6. Carpels 2- or 3-locular. N to C Europe, Siberia, the Himalayas, China, Japan, N to C America. – Polygonatum Mill. (71). Leaves lateral and pseudoterminal, distichous. Inflorescence axillary, racemose, umbel-like or 1-flowered. Flowers tubular to campanulate, urceolate, usually nodding. Tepals 6, connate, corona absent. Stamens 6, attached to tepals. Ovules 4–6 per locule. Fruit a dark purple to orange or red berry. Seeds pale brown. Temp. Eurasia and N America, diversity centers in the Himalayas, China, Japan.Subfam. Dracaenoideae Engl. (incl. Dracaenaceae Salisb., Sansevieriaceae Nakai) Trees, shrubs or climbers to geophytes with rhizomes, from 10 cm to 40 m tall. Stems often with leaf scars. Secondary growth via secondary thickening meristeme present. Leaves alternate, dis-tichous or helical, entire, ovate to ensiform, amplexicaul. Inflorescence terminal or axillary, racemose, capitate or paniculate. Flowers grouped together on articulated pedicels, white, flushed with green or purple, fragrant, nocturnal. Tepals 6, basally connate forming a tube; free tepals equal. Stamens 6, as long as tepals; anthers versatile. Ovary superior, 3-locular. Ovules 1 per locule. Septal nectaries present. Fruit a bright orange berry or “gymnosper-mous” (Sansevieria).
Chrysodracon (Jankalkski) P.-L.Lu & Morden (incl. Pleomele Salisb. p.p.) (6). Trees. Leaves helical, subcoriaceous. Inflorescence a terminal panicle subtended by leaf bract ½ to equaling panicle branch length. Flowers yellow, tubular, funnelform. Staminal filaments flattened. Hawaii Is. – Dracaena L. (incl. Pleomele Salisb. p.p.) (116). Trees (up to 40 m tall) or shrubs with woody trunks or stems. Leaves in tufts or rosettes. Inflorescence usu-
Fig. 4-101. Asparagales. Ruscaceae. 1–2. Dracaena draco. 1. Habit. 2. Detail of habit. 3–4. D. surcu-losa. 3. Inflorescence. 4. Detail of inflorescence. 5–6. Sansevieria spec. Detail of inflorescence. (Orig.; 1–6 phot. E. Fischer) (Scale bar 1: 5 m, 2: 1 m, 3: 5 cm, 4–6: 1 cm).
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Isotrema arborea (Linden) Eb.Fisch. comb. nov., p. 154Nolinoideae Eb.Fisch. & G.Mwachala subfam. nov., p. 431
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Sources of IllustrationsOrig.: Figs 3-6.–3-7., Figs 3-9.–3-12., Figs 3-15.–3.23., 4-2.–4-104. (Notice courtesy.)APG III. – Bot. J. Linn. Soc. 161: 105–121: Fig. 4-1.Farjon, A. A Handbook of the World’s Conifers, Brill, Leiden/Boston: Fig. 3-13. (Courtesy A. Farjon).Schweizerische Orchideenstiftung am Herbarium Jany Renz: Fig. 4-76.Strasburger, Lehrb. Bot., 35. Aufl., 2002. Spektrum, Akad. Verlag; Heidelberg, Berlin: Fig. 11-289, Fig.
11-207: Figs 3-1.–3-2. (Rights and Permissions by Springer Science+Business Media, Tiergartenstr. 17, 69121 Heidelberg, Germany.)
Syllabus der Pflanzenfamilien, 12. ed., I, 1954. Borntraeger, Berlin: Figs 3-3.–3-5., Fig. 3-8., Fig. 3-14.
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Index to Taxa (Subkingdom to genus level)Names in Roman type are accepted names. Names in italics are synonyms or invalid names. Bold page numbers are principal references, if there are more numbers. Numbers with an asterisk (*) refer to figures.
Part 4 of Engler’s Syllabus of Plant Families – Pinopsida (Gymnosperms) and Magnoliop-sida (Angiosperms) p.p.: Subclass Magnoliidae [Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Asparagales)] provides a thorough treatise of the world-wide morphological and molecular diversity of the Gymnosperms and the rst part of the Angiosperms [Magnoliidae: Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Asparagales).]
The description of the Gymnosperms, including the extinct diversity, is the rst synthesis of classical anatomical-morphological characters with modern molecular data, combined with the numerous new discoveries of fossils, especially from China, made during the last ten years.
The Angiosperms are the most diverse group of plants and form nearly 95% of the global vegetation from arctic tundra, resp. subantarctic vegetation formations, to tropical rainforests. There is actually no comprehensive survey covering all families and genera of angiosperms. En-gler’s Syllabus is an attempt to ll this gap by covering all angiosperms in two volumes arranged according to the most recent phylogenetic system of APG III (2009). In this rst volume (Part 4 of the Syllabus) all families and genera of Magnoliids are described. The monocotyledonous fami-lies are covered by the orders Acorales and Alismatales as well as all groups of Liliid orders and families (Petrosaviales, Dioscoreales, Pandanales, Liliales and Asparagales). The Orchidaceae are included with a fully revised and modern treatise, thus representing one of the two most spe-cies-rich families of plants. The remaining monocotyledonous groups (Arecales, Commelinales, Poales, Zingiberales and Dasypogonales) and the core eudicotyledons will be treated in Part 5 of the Syllabus of Plant Families. This up-to-date overview of the Pinopsida (Gymnosperms) and Magnoliopsida (Angiosperms) p.p. (Subclass Magnoliidae p.p.) will be of service as a valuable reference for a long time.
Engler’s Syllabus of Plant Families has since its rst publication in 1887 aimed to provide both the researcher, and particu larly the student with a concise survey of the plant kingdom as a whole, presenting all higher systematic units right down to families and genera of plants and fungi. In 1954, more than 60 years ago, the 12th edition of the well-known „Syllabus der P anzenfamilien“ (“Syllabus of Plant Families”), set a standard.
Now, the completely restructured and revised 13th edition of Engler’s Syllabus published in 5 parts and in English language for the rst time also considers molecular data, which have only recently become available in order to provide an up-to-date evolutionary and systematic overview of the plant groups treated.
In our “molecular times” there is a vitally important and growing need to preserve the knowledge of the entire range of diversity and biology of organisms for coming generations, as there is a decline in “classical” morphological and taxonomical expertise, especially for less popular (showy) groups of organisms.
Accordingly, the 13th edition of Syllabus of Plant Families synthesizes both modern data and classical expertise, serving to educate future experts who will maintain our knowledge of the full range of Earth’s biodiversity. Syllabus of Plant Families is a mandatory reference for students, experts and researchers from all elds of biological sciences, particularly botany.
Part 4Pinopsida (Gymnosperms), Magnoliopsida (Angiosperms) p.p.:Subclass Magnoliidae [Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Asparagales)]
Syllabus of Plant Families Wolfgang Frey (Editor)2015. XI, 495 pp., 127 colour plates, 1 tablehardcover, 25 x 17 cmISBN 978-3-443-01087-4 139.– €www.borntraeger-cramer.com/9783443010874
The Syllabus of Plant Families is a mandatory reference for students, experts and researchers from all elds of biological sciences, particularly botany.
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Part 4 of Engler’s Syllabus of Plant Families – Pinopsida (Gymnosperms) and Magnoliopsida (Angiosperms) p.p.: Subclass Magno-liidae [Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Asparagales)] provides a thorough treatise of the world-wide morpho-logical and molecular diversity of the Gymnosperms and the rst part of the Angiosperms [Magnoliidae: Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Asparagales).]
The description of the Gymnosperms includes the diversity of extinct species and is the rst synthesis of classical anatomical-morphological characters and modern molecular data, combined with the numerous new discoveries of fossils, especially from China, made during the last ten years.
Angiosperms form the most diverse group of plants and make up nearly 95% of the global vegetation from the arctic tundra, resp. subantarctic vegetation formations, to tropical rainforests. There is actually no comprehensive survey covering all families and genera of angiosperms. Engler’s Syllabus attempts to ll this gap by co-vering all angiosperms in two volumes arranged according to the most recent phylogenetic system of APG III (2009). In this rst volume (Part 4 of the Syllabus) all families and genera of Magno-liids are described. The monocotyledonous families are covered by the orders Acorales and Alismatales as well as all groups of Liliid orders and families (Petrosaviales, Dioscoreales, Panda-nales, Liliales and Asparagales). The Orchidaceae are included with a fully revised and modern treatise, thus representing one of the two most species-rich families of plants. The remaining mo-nocotyledonous groups (Arecales, Commelinales, Poales, Zingi-berales and Dasypogonales) and the core eudicotyledons will be treated in Part 5 of the Syllabus of Plant Families. This up-to-date overview of the Pinopsida (Gymnosperms) and Magnoliopsida (Angiosperms) p.p. (Subclass Magnoliidae p.p.) will be of service as a valuable reference for a long time. Sample page from Syllabus of Plant Families Part 4
4 Pinopsida (Gymnosperms), Magnoliopsida (Angiosperms) p.p. Syllabus of Plant Families
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