Zootaxa, Cnidaria, Octocorallia, Paragorgiidaedocs.niwa.co.nz/library/public/1877407194.pdf · Systematics of the bubblegum corals (Cnidaria: Octocorallia: Paragorgiidae) with description

ZOOTAXA

Systematics of the bubblegum corals (Cnidaria: Octocorallia: Paragorgiidae)

with description of new species from New Zealand and the Eastern Pacific

JUAN ARMANDO SÁNCHEZ

Magnolia PressAuckland, New Zealand

1014

JUAN ARMANDO SÁNCHEZ

Systematics of the bubblegum corals (Cnidaria: Octocorallia: Paragorgiidae) with description of new species from New Zealand and the Eastern Pacific(Zootaxa 1014)

72 pp.; 30 cm.

5 July 2005

ISBN 1-877407-18-6 (paperback)

ISBN 1-877407-19-4 (Online edition)

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ISSN 1175-5326 (Print edition)

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1014

Accepted by P. Alderslade: 26 May 2005; published: 5 Jul. 2005 3

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2005 Magnolia Press

Zootaxa 1014: 1–72 (2005) www.mapress.com/zootaxa/

Systematics of the bubblegum corals (Cnidaria: Octocorallia: Par-agorgiidae) with description of new species from New Zealand and the Eastern Pacific

JUAN ARMANDO SÁNCHEZNational Institute of Water and Atmospheric Research-NIWA, Wellington, New Zealandand Departamento de Ciencias Biológicas, Universidad de los Andes, Carrera 1E No 18A – 10, P.O.Box 4976, Bogotá, Colombia. Email: [email protected]

Table of contents

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Morphological characters and phylogenetic relationships . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Key to the species of Paragorgiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14Family PARAGORGIIDAE Kükenthal, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

Genus Paragorgia Milne Edwards & Haime, 1857 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15Paragorgia arborea (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15Paragorgia johnsoni Gray, 1862 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21Paragorgia splendens Thomson & Henderson, 1906 . . . . . . . . . . . . . . . . . . . . . . . . 23Paragorgia regalis Nutting, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25Paragorgia coralloides Bayer, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29Paragorgia alisonae sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31Paragorgia kaupeka sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37Paragorgia maunga sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41Paragorgia aotearoa sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44Paragorgia wahine sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47Paragorgia whero sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49Paragorgia tapachtli sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51Paragorgia yutlinux sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53Paragorgia stephencairnsi sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57

Genus Sibogagorgia Styasny, 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60Sibogagorgia weberi Stiasny, 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60Sibogagorgia tautahi sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62Sibogagorgia dennisgordoni sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64

SÁNCHEZ4 © 2005 Magnolia Press

1014ZOOTAXA Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67

Acknowledgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

Abstract

Bubblegum corals, Paragorgiidae, are among the largest and most ecologically important benthicsessile deep-water organisms harboring hundreds of associated. However, no recent reviews of theirdiversity and systematics are yet available, despite the recent increase in the sampling and fishingof deep-water habitats. This study covers 17 Paragorgiidae species. There were only five previouslyknown species for Paragorgia (P. arborea [Linnaeus], P. johnsoni Gray, P. splendens Thomson &Henderson, P. regalis Nutting [=dendroides Bayer], and P. coralloides Bayer) and just one ofSibogagorgia (S. weberi Stiasny). Eleven new species are described here comprising 9 Paragorgiaspp. (P. alisonae, P. kaupeka, P. maunga, P. aotearoa, P. wahine, P. whero, P. yutlinux, P.stephencairnsi, and P. tapachtli) and 2 Sibogagorgia spp. (S. tautahi and S. dennisgordoni). Thisstudy also uncovered two areas of endemism for bubblegum corals corresponding to New Zealandand the Eastern Pacific (Mexico to Canada). New Zealand has 6 likely endemic species of Paragor-gia (P. alisonae, P. kaupeka, P. maunga, P. aotearoa, P. whero, and P. wahine) and the two new spe-cies of Sibogagorgia, whereas P. yutlinux, P. stephencairnsi, and P. tapachtli were collected in theEastern Pacific. There seem to be a few trans-Pacific species such as P. regalis, and likewise in theAtlantic with P. johnsoni, but it is clear that no other species is as cosmopolitan as P. arborea withdiscontinuous but bi-polar distribution. There are cases of morphological sister species such as P.johnsoni and P. aotearoa that correspond to the Atlantic and Pacific respectively, but the phyloge-netic relationships of the remaining species indicate that most paragorgiid diversity and speciationtook place in the Indo-Pacific region, as suggested by a number of sympatric species. Surface scler-ites, radiates, exhibit a great deal of variation under the Scanning Electron Microscope (SEM), pro-viding a number of characters for phylogenetic reconstruction, including three kinds of radialornamentation and several types of surfaces and sub-ornamentation at the ultrastructure level. Thethree most parsimonious trees of equal length, using morphological characters, showed P. arboreaas basal to the rest of the Paragorgia species (using Sibogagorgia as the outgroup), which weredivided in two clades. One clade includes the species with asymmetrical surface sclerites with someradial ornaments larger or different than others have ([P. maunga -[P. coralloides-P. tapachtli-P.regalis-P. kaupeka]]). In this clade, P. maunga conserved the basal position in the most parsimoni-ous trees whereas relationships among the other species were not consensual. The other clade com-prised species with symmetrical surface sclerites ([[P. splendens-P. wahine] P. alisonae-[P.yutlinux-P. stephencairnsi]-P. johnsoni-P. aotearoa]). P. splendens-P. wahine-P. whero and P. yut-linux-P. stephencairnsi maintained their sister relationships respectively in all most-parsimonioustrees but no consensual relationships with respect to and among the other species of the clade. Com-plete descriptions of described and new species using SEM, species comparisons, character states,and a species key are also provided in this paper.

Key words: Paragorgiidae; Paragorgia; Sibogagorgia; New Zealand; seamounts; Eastern Pacific;octocoral; phylogeny; Octocorallia; deep-water coral; bubblegum corals; Coelenterata; Cnidaria

© 2005 Magnolia Press 5PARAGORGIIDAE

1014ZOOTAXAIntroduction

Deep-water octocorals are among the largest sessile benthic invertebrates on continentalshelves and seamounts, where most fishery activities occur, providing habitat for associ-ated fish and invertebrates (e.g., Krieger 1993; Koslow et al. 2001; Heifetz 2002; Krieger& Wing 2002). Samples from just seven colonies of the bubblegum coral, Paragorgiaarborea Linnaeus 1758, have been recorded harboring over 1300 crustacean individualsfrom 16 different associated-species (Buhl-Mortensen & Mortensen 2004). Nonetheless,in such fragile environments, an extractive activity such as trawling, which disturbs theseabed, have started to endanger not only fishery resources but also important habitat-forming organisms such as corals (e.g., Watling & Norse 1998). Unfortunately, some ofthe largest habitat-forming deep-water organisms have not been formally described (e.g.,Sánchez & Cairns 2004), which prevents us from constructing a proper account of theirdiversity and carrying out further conservation actions. Consequently, it is a priority todescribe important long-lived habitat-forming organisms such as the bubblegum corals,the Paragorgiidae, which attain the largest sizes known among extant octocorals.

The Paragorgiidae are dimorphic octocorals with large polyps (autozooids), usuallyconcentrated in dispersed clusters, and numerous tiny siphonozooids (presumably pump-ing and reproductive polyps) generally occurring throughout the colony surface. The col-ony appearance of robust branches with soft clumps of polyps awarded paragorgiids thecommon name bubblegum corals. Paragorgiids are unusual octocorals because, despiteforming enormous tree-like structures, they do not contain a corneous or calcareous axialskeleton like most branching gorgonians. Branches of bubblegum corals are rigid owing tothe accumulation and near fusion of microscopic calcitic sclerites (Bayer 1993). Paragor-gia was originally the only paragorgiid genus. Bayer (1956a) included Sibogagorgia in thefamily, despite Verseveldt (1942) having created a separate family Sibogagorgiidae.Sibogagorgia has many characteristics similar to Paragorgia but lacks polyp sclerites andmedullar canals while having a reticulate network of boundary canals in the outer medulla/subsurface. However, polyp polymorphism and other sclerite types and forms support anobvious relationship between Sibogagorgia and Paragorgia. Since the aim of systematicsis to understand the relationships among species, it should be clear that Paragorgia andSibogagorgia are members of a monophyletic group, the Paragorgiidae, as will be illus-trated in detail in this paper.

The closest paragorgiid relatives, according to both molecular and ultrastructural data,are the precious corals of the family Coralliidae (e.g., Sánchez et al. 2003). They have thesame sclerite types, identical microcrystals forming the sclerites, similar branching pat-terns, and various degrees of sclerite fusion, with the extreme example being Coralliumspp. (Bayer 1992; Sánchez et al. 2003). Both Corallium and Paragorgia have been classi-fied in the suborder Scleraxonia; gorgonians (Gorgonacea) with sclerites in the colonyaxis. However, molecular phylogenies have shown that Scleraxonia is largely polyphyleticand Corallium and Paragorgia belong to a clade together with Isididae and Primnoidae


1014ZOOTAXA even though little morphological affinity among these families is observable (e.g., Sánchez

et al. 2003). It is generally described in the taxonomic literature for Corallium that the axisis composed of fused sclerites. However, studies on the physiology of calcium depositionand skeletogenesis have shown that the axis in Corallium results from a secretion by akind of epithelium surrounding the axis (see review in Allemand 1993), which supportsthe affinity with other non-sclerite-derived hard-axis gorgonians such as Isididae andPrimnoidae. Nonetheless, species such as Paragorgia coralloides Bayer, 1993 have verysimilar sclerites to those of Corallium spp. (Bayer 1993) but lack a rigid calcareous axis,and some New Zealand species of Corallium (unpublished) have sclerites similar to thoseof P. splendens Thomson & Henderson (1906). Consequently, it could be possible thatParagorgiidae and Coralliidae are not entirely monophyletic. Whether or not the Coralli-idae is just a derived branch of the Paragorgiidae requires further study with the aid ofindependent characters.

The aim of this paper is to clarify systematic relationships among the New ZealandParagorgiidae using morphological criteria. Owing to the complexity involved with octoc-oral taxonomy, and particularly the Paragorgiidae, the New Zealand fauna was reviewed incomparison to all described species and some undescribed material from the EasternPacific (Fig. 1). Thus, this paper provides the most current systematic revision of the Para-gorgiidae worldwide, including a systematic analysis of morphological characters, a spe-cies key, and diagnosis/description of both described and new species from New Zealandand the Eastern Pacific (Canada, USA, and Mexico).

FIGURE 1. Geographical locations of the known Paragorgiidae species (see Fig. 2 for NewZealand detail).


1014ZOOTAXA

FIGURE 2. Map of New Zealand and overall underwater landscape showing the locations and dis-tribution of Paragorgiidae species.

Material and Methods

The study of the Paragorgiidae was based primarily on material collected throughout theNew Zealand Economic Exclusive Zone (EEZ), by multiple cruises during several years(Fig. 2) and deposited in the NIWA Invertebrate Collection (National Institute of Water &Atmospheric Research), Wellington. Voucher specimens and types from Eastern Pacificsamples were already deposited at the National Museum of Natural History, SmithsonianInstitution, Washington, D.C., USA (USNM). Preliminary identification of the Paragorgiaspecies were based on taxonomic works by Bayer (1956b; 1964; 1993) and Grasshoff(1979), which include revisions of Paragorgia worldwide. Type material from describedspecies was reviewed when possible. Samples were examined through dissection of scler-ite layers, digestion of the organic matter in sodium hypochlorite, and observations under


1014ZOOTAXA optical compound microscope (200 x). For some sclerite types, 10 haphazardly chosen

individual sclerites per species were measured using a micrometer and compound micro-scope. Detailed images from sclerite ultrastructure were obtained with Scanning ElectronMicroscopy-SEM following the methods from Sánchez & Cairns (2004), for stereo-pairimages of some sclerites, using a Cambridge Instruments-Leo 440 at 10 kv (Gracefield,Industrial Research Ltd, Lower Hutt, New Zealand) and optimum magnification for eachsclerite type.

Most colonial features are highly polymorphic and not possible to code consistently ineach species. Informative characters pertain primarily to the microscopic calcite sclerites,which exhibit a great deal of variation under the SEM. Paragorgiids have three mainhomologous sclerite types, here referred to as surface (or surface cortex), autozooid tenta-cle (absent in Sibogagorgia spp.), and medullar sclerites, corresponding to the namedstructures. There is a wide variation in sclerite forms in paragorgiids, resulting in a contin-uum between the surface (including the autozooid calyx), with the smaller forms, and theinner medulla which has larger forms. It is commonplace in paragorgiids to have multiplepolymorphic forms in the subsurface/outer medulla, though not in all species. Therefore,in order to make homologous comparisons among species, sclerites from the two extremesof the continuum were carefully dissected and prepared for SEM. Otherwise, it wouldhave been impossible to construct a key based on these characters as well as phylogeneticrelationships, particularly since most of the characters are based on the surface sclerites.Nonetheless, in each species description a note on the variation of the different scleriteform is included. Following the taxonomical convention for Paragorgia (e.g., Bayer1956b; 1961; 1964; 1993; Grasshoff 1979), sclerites are referred to as radiates (surfacecortex), blunt rods or sclerites (autozooid polyps), and spindles (medulla), but it is impor-tant to note that those character names may not be considered homologous for the mem-bers of other families where these names have been used (e.g., Plexauridae: Sánchez 2001;Sánchez & Cairns 2004).

Morphological characters include the topographical position of sclerites in the polyp,tissue layers in relation to the colonial axis, and special similarity as criteria for homologyamong species (e.g., Sánchez 2001) as well as outgroup analyses for character polarization(e.g., Kitching et al. 1998). Since most of the species in this study belong to Paragorgia,and there is not yet clarification concerning a closely related outgroup for Paragorgiidae,Sibogagorgia was used as the outgroup. The best optimal phylogenetic trees weresearched with the branch-and-bound algorithm using maximum parsimony criterion inPAUP* (Swofford 2002) and the tree editor Winclada for character mapping (Nixon1999). To examine the effect of homoplasy in the data set, the Permutation Tail Probability(PTP) was also determined using PAUP* (Fu & Murphy 1999) in comparison to the resultsfrom 100 random trees replicates.


1014ZOOTAXAResults

Morphological characters and phylogenetic relationships

Table 1 presents the 12 informative morphological characters examined among Paragorgi-idae species, 8 of which were possible to code discretely for phylogenetic analyses. Onlytwo colonial characters, terminal branch width and cortex color, were included in phyloge-netic analyses. Surface radiates show a great deal of variation, including three kinds ofradial ornamentation (e.g., Fig. 3) and several types of surfaces and sub-ornamentation atthe ultrastructure level (Table 1). (All characters are discussed in detail in every speciesdescription). There is also considerable variation in the size of surface sclerites but thisvariation is not suitable for discrete coding owing to overlap among paragorgiid species(Fig. 4A). In order to incorporate part of the variation in the size of surface sclerites, thelength/width ratio (Fig. 4B) was coded discretely for inclusion in the phylogenetic analy-ses. This metric character included information on the form of the sclerite from nearlyspherical forms (e.g., length/width ratio 1.3) to more spindle-like forms (e.g., >1.65) aswell as forms in between (>1.5 and < 1.65). This ratio also provides information on howmuch longer the sclerite is in relation to its width. The other two sclerite types, surface andmedullar, do not present any straightforward character states within Paragorgia orSibogagorgia but provide a clear differentiation between them as well as valuable autapo-morphies for species descriptions and comparisons.

FIGURE 3. Major types of surface cortex radiate sclerites: A–B, 6-radiates; C, 7-radiate; D, 8-radi-ate.


1014ZOOTAXA

FIGURE 4. A. Box plots from the distribution and variance of surface cortex (n=10) sclerites ofthe examined specimens. The median line is inside the 25th and 75th percentiles with external errorbars at the 10th and 90th percentiles. B, Scatter plot of the surface cortex sclerites length vs width(different symbols indicate the different specimens).

Despite the low number of informative morphological characters among paragorgiidsclerites, as in other octocorals (e.g., Sánchez 2001), eight characters provided three par-tially resolved most parsimonious trees of Length = 19 (Consistency index=0.78, Reten-tion Index=0.88, and Homoplasy index=0.21) using Sibogagorgia as the outgroup. In spiteof this, only three nodes varied among the two most parsimonious trees (Fig. 5A–B). Theunpermuted most-parsimonious trees were significantly shorter than 100 random permuta-tion replicates averaging a length of 44 steps (2.37 SD, Length >33, PTP: P<0.05) signify-ing that the phylogenetic result scores and topologies could not be matched by randomphylogenies of Paragorgiidae. All the most parsimonious trees had a clear separationbetween Paragorgia and Sibogagorgia species, including a divergence of three characters(characters/states: 2/2, 7/1, 8/1: Fig. 5A–B). All the trees showed Paragorgia arboreabasal to the rest of Paragorgia species, which were divided into two clades. One cladeincludes the species with asymmetrical surface sclerites with some ornaments larger or dif-ferent than others ([P. maunga -[[P. coralloides-P. tapachtli] [P. regalis-P. kaupeka]]]: Fig.5). That clade conserved one topology among the most parsimonious trees (Fig. 5). Theother clade had species with symmetrical surface sclerites ([[P. splendens-P. whero-P.wahine][P. alisonae-[P. yutlinux-P. stephencairnsi]-P. johnsoni-P. aotearoa]]: Fig. 5). OnlyP. yutlinux-P. stephencairnsi maintained their sister relationship respectively in all mostparsimonious trees but no consensual relationships with the other species of the clade


1014ZOOTAXAoccurred (Fig. 5). Additional putative relationships and comparisons among paragorgiids,

based on special similarity of certain characters, are discussed following every speciesdescription below. Nevertheless, one of the most parsimonious trees (Fig. 5B), as well asthe character descriptions from Table 1, provided a good basis for a key to the new andpreviously described species of Paragorgiidae which follow.

FIGURE 5. Cladogram of Paragorgiidae species using morphological characters corresponding to

two of the partially resolved most parsimonious trees (A–B); Adams consensus tree (C) and strict

consensus (D). Hash marks are mapped characters/states (above/below respectively) from Table 1

and discontinuous states only (open bars) correspond to homoplasy.


1014ZOOTAXA TABLE 1. Morphological characters and states for the known species of Paragorgiidae. Numbers

in the characters column relate to the characters in Fig. 5 and in the matrix deposited in TreeBase(Acc. No. S1284 & M2241).

Characters States Paragorgia

arbo

rea

john

soni

sple

nden

s

cora

lloid

es

rega

lis

alis

onae

kaup

eka

mau

nga

aote

aroa

yutli

nux

step

henc

airn

si

wah

ine

Terminal branch width (1)

(0) < 4mm (1) > 5 mm 1 0 0 0 0 0 0 0 0 0 0 0

Coenenchyme colour (2)

(0) Colourless, (1) pink, (2) red, (3) Colourless and dark (purple) autozooid apertures

2 2 1 1 1 2 1 2 2 3 3 1

Surface sclerite form (3)

(0) Symmetrical (all rays equal), (1) Asym-metrical (some larger or differentiated rays present)

0 0 0 1 1 0 1 1 0 0 0 0

Surface sclerite rays (=warts) (4)

(0) Globular & fused (1) smooth (unfused), (2) Grooved (3) Lobu-lated (4) > 5 lobes

2 1 4 0 2 4 2 2 1 3 3 4

Surface sclerite length Average (mm) 0.042 0.055 0.068 0.05 0.07 0.079 0.049 0.07 0.076 0.05 0.065 0.071

Surface sclerite width Average (mm) 0.033 0.035 0.038 0.03 0.043 0.052 0.028 0.04 0.046 0.034 0.045 0.04

Surface sclerite length/width

Ratio 1.27 1.57 1.79 1.68 1.61 1.51 1.75 1.75 1.65 1.47 1.45 1.71

Surface sclerite length/width (coded) (5)

(0) <1.3, (1) 1.5<X<1.65, (2) >1.66

0 1 2 2 2 1 2 2 1 1 1 2

Surface sclerite orna-mentation (dominant forms) (6)

(0) 6-radiates, (1) 7-radiates, (2) 8-radiates

0 2 2 2 1 2 1 2 2 0 1 2

Medulla sclerite size Continuous (max, mm)

0.5 0.38 0.7 0.15 0.5 0.4 0.4 0.35 0.33 0.25 0.3 0.3

Medulla spindles (7) (0) Smooth (few or no ornamentation), (1) Ornate

1 1 1 1 1 1 1 1 1 1 1 1

Autozooid tentacle sclerites (8)

(0) Absent, (1) Ornate blunt sclerites

1 1 1 1 1 1 1 1 1 1 1 1


1014ZOOTAXATABLE 1. Continued.

Characters States Paragorgia Sibogagorgia

step

henc

airn

si

wah

ine

whe

ro

tapa

chtli

web

eri

taut

ahi

denn

isgo

rdon

i

Terminal branch width (1)

(0) < 4mm (1) > 5 mm 0 0 0 0 1 1 1

Coenenchyme colour (2) (0) Colourless, (1) pink, (2) red, (3) Colour-less and dark (purple) autozooid apertures

3 1 1 1 0 0 0

Surface sclerite form (3) (0) Symmetrical (all rays equal), (1) Asym-metrical (some larger or differentiated rays present)

0 0 0 1 0 0 0

Surface sclerite rays (=warts) (4)

(0) Globular & fused (1) smooth (unfused), (2) Grooved (3) Lobulated (4) > 5 lobes

3 4 4 0 3 3 2

Surface sclerite length Average (mm) 0.065 0.071 0.089 0.043 0.064 0.08 0.069

Surface sclerite width Average (mm) 0.045 0.04 0.052 0.026 0.039 0.047 0.045

Surface sclerite length/width

Ratio 1.45 1.71 1.71 1.63 1.64 1.71 1.53

Surface sclerite length/width (coded) (5)

(0) <1.3, (1) 1.5<X<1.65, (2) >1.66 1 2 2 2 1 2 2

Surface sclerite orna-mentation (dominant forms) (6)

(0) 6-radiates, (1) 7-radiates, (2) 8-radiates 1 2 2 2 2 2 2

Medulla sclerite size Continuous (max, mm) 0.3 0.3 0.5 0.3 0.3 0.25 0.4

Medulla spindles (7) (0) Smooth (few or no ornamentation), (1) Ornate

1 1 1 1 0 0 0

Autozooid tentacle scler-ites (8)

(0) Absent, (1) Ornate blunt sclerites 1 1 1 1 0 0 0


1014ZOOTAXA Key to the species of Paragorgiidae

1a. Medulla perforated by 3 or more large canals in terminal branches; autozooid tenta-cles with sclerites.................................................................................Paragorgia (4)

1b. Medulla not perforated by large canals in terminal branches; autozooid tentaclesdevoid of sclerites.............................................................................Sibogagorgia (2)

2a. Medulla sclerites ornate throughout..................................................S. weberi Stiasny2b. Medulla sclerites with ornaments only at the tips (bare body) ............................... (3)3a. Medulla sclerites thick and blunt; surface radiate sclerites with lobulated ornaments

......................................................................................................... S. tautahi sp. nov.3b. Medulla sclerites thin and long; surface radiate sclerites with smooth and grooved

ornaments .............................................................................S. dennisgordoni sp. nov.4a. Terminal branches (excluding autozooid clumps) > 5 mm in diameter; surface radi-

ate sclerites with subdivided and grooved ornaments..................P. arborea Linnaeus4b. Terminal branches (excluding autozooid clumps) < 5 mm in diameter; surface radi-

ate sclerites smooth, globular, irregular, subradiate, and/or lobate ornaments ....... (5)5a. Surface sclerites asymmetrical (some rays larger or differentiated) ....................... (6)5b. Surface sclerites symmetrical (all rays equal)....................................................... (10)6a. Pink with a thick cortex .......................................................................................... (7)6b. Red, fleshy with a thin cortex........................................................P. maunga sp. nov.7a. Surface sclerites predominantly smooth to lobulated; 7-radiates ........................... (9)7b. Surface sclerites with globular and fused rays; 8-radiates...................................... (8)8a. Some radial ornaments subdivided in non-globular lobes..........P. coralloides Bayer.8b. All radiates globular with few or no subdivisions........................P. tapachtli sp. nov. 9a. Surface sclerites averaging 0.05 mm in length..............................P. kaupeka sp. nov.9b. Surface sclerites averaging 0.065 mm in length ..............................P. regalis Nutting10a. Surface sclerites about 1.7 times longer than wider; pink cortex ......................... (13)10b. Surface sclerites between 1.5 and 1.65 longer than wider; red, or polychromatic cortex

................................................................................................................................(11)11a. Slim terminal branches (up to 2 mm in diameter); autozooid polyps fully retracted;

tightly closed apertures ......................................................................................... (12) 11b. Terminal branches 3–4 mm in diameter; autozooid polyps exert; wide-open apertures

........................................................................................................... P. whero sp. nov. 12a. Terminal branches 1 mm; gently curved branch tips ..................................................

............................................................................P. splendens Thomson & Henderson 12b. Terminal branches 2 mm; straight but irregular branch tips........... P. wahine sp. nov.13a. Surface sclerites with ornaments with >5 lobes............................ P. alisonae sp. nov.13b. Surface sclerites with ornaments with few (<5) or no lobes................................. (14)14a. Surface sclerites with smooth and rounded rays; red cortex................................. (15)14b. Surface sclerites with lobulated rays; white with pink/purple autozooid apertures …

...............................................................................................................................(16)


1014ZOOTAXA15a. Surface sclerites averaging 0.05 mm in length................................. P. johnsoni Gray

15b. Surface sclerites averaging 0.07 mm in length.............................P. aotearoa sp. nov.16a. Surface sclerites mostly 6-radiates.................................................P. yutlinux sp. nov.16b. Surface sclerites mostly 7-radiates.......................................P. stephencairnsi sp. nov.

Systematic account and species descriptions

Family PARAGORGIIDAE Kükenthal, 1916

Diagnostic characters. Dimorphic octocorals, reproductive siphonozooids and feedingautozooids, without axial skeletal structures other than a medulla formed by unfused scler-ites. Medulla surrounded by a cortex containing the gastric cavities of autozooids andsiphonozooids. Profusely branched colonies.

Genus Paragorgia Milne Edwards & Haime, 1857

Diagnostic characters. Paragorgiidae with axial medulla of unfused spindles up to 0.6mm in length; other sclerites less than 0.15 mm in length. Different kinds of sclerites in theautozooid polyp tentacles, cortex surface, medulla, and subsurface/outer medulla. Surfacesclerites are 6 to 8 radiates, always less than 0.1 mm in length, with globular, smooth,grooved or lobulated ornaments. Medulla sclerites are ornate spindles usually less than 0.8mm in length. Intermediate forms between radiates and spindles in the subsurface/outermedulla. Scleritic medulla perforated at terminal branches with 3–7 main canals, andnumerous smaller canals occurring both in the medulla and subsurface/outer medulla (seedetails in Verseveldt, 1940). Autozooid polyp tentacles with distinctive blunt, stubby rodsor ovals usually less than 0.1 mm.

Paragorgia arborea (Linnaeus, 1758)(Figs. 6–9)

Alcyonium arboreum Linnaeus 1758: 803. Paragorgia nodosa Koren & Danielsen 1883 (sensu Bayer 1956: 70).Paragorgia pacifica Verrill 1922: G16.Paragorgia arborea: Broch 1912: 6; Grasshoff 1979: 117 (and references therein). Material examined: NIWA 3308, 33°55.6’S–167°55.1’E, 1225 m, (Waipori, bottomtrawling, Z10987), 23 January 2002, New Zealand; NIWA 3309, (J217), 33°55.6’S–

167°54.4’E, 955 m, (Waipori 1621/011, Z11009) 8 June 2002; NIWA Z10920 (J208)/Z10956 (J212), (AEX 0101/080), New Zealand EEZ, exact locality unknown (probably


1014ZOOTAXA near: 42°47’S–179°32’E, 1077 m), (Kap Farval), 17 October 2001; NIWA 3310, Z9862

(J219), 44°45.0S–174°50.0–49.0’E, 687–940 m, (Kap Farval 1278/102), 21 October 1999,New Zealand; NIWA 3311, X700 (J221), 35° 50.447’–49.900’S–177° 54.497–54.644’E,“Southern Havre trough” (dredge), 1525–1798 m, 12 February 1996, New Zealand;USNM 98045, 52° 17’–52° 22’S–160° 40’–160° 34’E, 659–798 m, (Eltanin R/V 1414,USAP 16, U. Southern California, trawl-blake), 9 February 1965, New Zealand; USNM52433, 54° 19’N–159° 40’ W, (Albatross R/V 3338), 1143 m, 28 August 1890, Alaska,USA; USNM 1014919, Davison seamount, 1313 m, California 2003, USA.

Diagnostic characters. Surface cortex (including calyx surface) containing small,ornate, radiate sclerites (6, 7, and 8 rays); predominantly 6 radiates with distinctive, laby-rinth-like, groove markings in the radiate projections or ornamentations (Figs. 7C–D and8C–D).

FIGURE 6. Colonies of Paragorgia arborea (Linnaeus): A–B, NIWA 3308; C–D, NIWA Z10920.Calliper open 2 cm.


1014ZOOTAXA

FIGURE 7. Paragorgia arborea (Linnaeus) (NIWA Z10920): A–B, stereo pairs of the scleritesfrom the polyps (scale 10 µm); C–D, radiate sclerites (stereo pairs) from the colony surface (scales10 µm); E, sclerites from the colony medulla (scales 100 µm); F, additional sclerites from the col-ony surface and intermediates towards the medulla (scales 10 µm).

Description. Robust tree-like colonies up to several meters in height with bubble-likeconcentrations of autozooids (Fig. 6). Complete specimens exhibiting dense and regularaccumulations of autozooid nodules or bulbs on distal or lateral branches, whereas the dis-tal main stem and branches are usually without nodules (Fig. 6A). Inter-nodular surfacewith numerous and uniformly distributed tiny siphonozooid apertures giving the colony agranular texture (Fig. 6D). Medulla perforated by 5–7 main stem canals in terminalbranches, surrounded by both red and colorless spindles; outer medulla with colorlesssclerites and numerous smaller canals. Polyps completely retracted and enclosed withinthe calicular (small conical protuberance of the nodule) cortex. Tentacle sclerites are blunt,stubby ovals up to 0.1 mm (Figs. 7A–B, 8A–B), commonly found in several Paragorgiaspp. (Bayer, 1993), but some of them pointed like a spindle (Fig. 8A). They have conicalornaments usually arranged bipolarly with a smooth neck, and at the surface exhibit a


1014ZOOTAXA granular appearance due to microcrystal calcite tips. Radiate sclerites (6-radiate) from the

surface are small, averaging 0.043 mm length in two different specimens, exhibiting lowvariation (±0.004 SD, n=10, NIWA 3308; ±0.002 SD, n=10, NIWA Z10920). Radiates 1.3times longer than wide, averaging 0.03 mm in width (±0.001 SD, n=10, NIWA 3308;±0.002 SD, n=10, Z10920) also occur. The rays are ornamented with distinctive markings(see diagnostic character) and the surface between is smooth. Subsurface rich in diverseradiates and intermediate forms between radiate and spindle attaining larger sizes (Figs.7F, 8E). Medulla sclerites long, slim spindles up to 0.5 mm length with irregular projec-tions, some quite prominent (up to 0.02 mm) and bifurcated (Figs. 7E, 8F). Occasionallyslightly curved, intermediate forms (short and long ornamented spindles) present betweenthe medulla and subsurface (see Grasshoff, 1979).

FIGURE 8. Paragorgia arborea (Linnaeus) (NIWA 3308): A, stereo pair of a sclerite from the pol-yps (scale 10 µm); B, additional sclerites from the polyps (scales 20 µm); C–E, radiate sclerites (C–D are stereo pairs) from the colony surface (scales 10 µm); F, sclerites from the colony medulla(scales 100 µm); F, additional sclerites from the colony surface and intermediates towards themedulla (scales 10 µm).


1014ZOOTAXAMorphological variation. Whereas sclerites and ultrastructure variation has been usu-

ally low among many specimens and geographical localities (e.g., Grasshoff, 1979), colo-nial features seem to have a great deal of variation. One New Zealand specimen (NIWAZ9566) had unusually large siphonozooids and autozooids with eight grooved notches inthe aperture not present in the rest of the material, but no scleritic differences were found.Most New Zealand specimens were red except a few paler (pink) specimens. The size ofthe autozooid nodules was another factor of variation. Some specimens had nodules of lessthan 20 mm in diameter (e.g., Fig. 6A–B) and some had up to 30 mm (Fig. 6C–D), the lat-ter corresponding to the largest specimens from the NIWA collection. However, only onespecimen of this morphotype is fairly complete and no correlation with colony size can bemade. Similar variation has been recorded in northern latitudes including the presence ofwhite individuals (Tendal 1992), which have not been found in New Zealand.

Distribution. P. arborea has one of the most intriguing distributions among octocorals(e.g., Fig. 1), it has been abundantly collected and observed towards the two poles in theNorth Atlantic and in sub-Antarctic waters (Broch 1912) but no intermediate populationshave been found so far. Tendal (1992) provided a complete compilation of P. arborearecords in the North Atlantic concluding that the species is usually distributed between200 and 1330 m in depth and a temperature of 4–8 °C. In the Southern Hemisphere it hasbeen reported from the Falkland Islands, on the Patagonian shelf. New Zealand records(e.g., 1525 m) are the deepest records for this species.

Species comparisons. Broch (1957) reported specimens of P. arborea for the firsttime in the Southern hemisphere. Grasshoff (1979) reviewed material, including someSEM analyses, from boreal and austral latitudes concluding that they belong to the samespecies. The material collected and studied from New Zealand waters presented the samediagnostic characters and size variation presented by Grasshoff’s (1979) SEMs. The diag-nostic characters of the surface radiate sclerites of P. arborea are not similar to any otherParagorgia species. Other scleritic characteristics are very similar to other Paragorgiaspp. such as the polyp (tentacles) and medulla sclerites, but the diversity of forms in thesubsurface is richer in P. arborea than in any of the studied species. However, it is stillunknown if the boreal and austral populations of P. arborea are indeed the same inter-breeding population and if there are stepping-stone populations in the tropics; furthergenetic testing would help to solve this problem. An important variation within P. arborea“populations” is present in the specimens from Alaska in the North Pacific, correspondingto P. pacifica Verrill (1922). The sclerites from a specimen from Alaska have the samediagnostic characters including the particular 6-radiates with grooved ornaments (Fig. 9D)but the sclerites from the medulla seem to be reduced in size and ornamentation withrespect to the New Zealand P. arborea (Figs. 7F, 8F vs 9H). The surface sclerites (6-radi-ates) were also found to be smaller than the other Paragorgia species examined (Fig. 3).Paragorgia pacifica was described on the grounds of colony form and as Verrill (1922)stated, “there may be doubt whether P. pacifica is not a variety of P. arborea”, because it


1014ZOOTAXA was described without sclerite examination. Nevertheless, the North Pacific populations of

P. arborea seem to be the most derived morphologically, but a more comprehensive revi-sion including type material and genetics is needed before reaching conclusions on P. paci-fica and the differentiation of south vs north P. arborea populations.

FIGURE 9. Paragorgia arborea (Linnaeus) (=pacifica Verrill) (USNM 52433): A–B, stereo pairsof the sclerites from the autozooid polyps (scale 10 µm); C, additional sclerites from the autozooidpolyps (scale 10 µm); D–F, radiate sclerites (stereo pairs) from the colony surface (scales 10 µm);G, additional sclerites from the colony surface and intermediates towards the medulla (scales 10µm); H, sclerites from the colony medulla (scales 100 µm).


1014ZOOTAXAParagorgia johnsoni Gray, 1862

(Fig. 10)

Paragorgia johnsoni Gray, 1862: 125; Grasshoff, 1979: 430 (and references therein).Paragorgia boschmai Bayer, 1964: 527.

Material examined: USNM 73767, 27° 05’ N–79°40’ W, 608 m, 21 Feb 1984 (AlvinDSR/V 1335), Bahamas; USNM 100898, 39°52.32’ N–067°25.19’ W, (col. M. Turnipseed,Delaware II R/V), USA.

Diagnostic characters. Polyps (autozooids) with blunt spindles with irregular orna-mentation. Surface cortex sclerites, including the autozooid aperture, are 7- and 8-radiatesclerites up to 0.06 mm averaging 0.05 mm in length.

Description. Profusely branched colonies up to 900 mm in length (Grasshoff 1979:Fig. 1) with terminal branches down to 2–4 mm in diameter (Grasshoff 1979: Figs. 2–3).Numerous short, lateral branches arise from the main branches with an irregular but clav-ate appearance (Grasshoff 1979: Fig. 1). Autozooid and siphonozooid polyps locatedtowards one side of the colony (Grasshoff 1979: Figs. 2–3). Autozooids clustered in dis-tinct nodules (Bayer 1964: 527). Autozooid polyp tentacles with blunt spindles, slightlythicker than in most Paragorgia spp., up to 0.08 mm in length, irregularly ornate with con-ical rays (Fig. 10A–B). Surface cortex, including autozooid aperture, with 6-, 7- and 8-radiate (predominant 8-radiates) sclerites up to 0.06 mm averaging 0.05 mm in length(±0.003 SD, n=10) (Fig. 10C–F). Rays nearly symmetrical with smooth surfaces. Surfaceradiates about 1.57 times longer than wider, averaging 0.03 mm in width (±0.002 SD,n=10). Medulla with ornate (oftentimes ramified or forked) spindles up to 0.38 mm inlength (Fig. 10G; see also Bayer 1964: 531) with smaller forms in the subsurface/outermedulla (Fig. 10H).

Morphological variation. P. johnsoni exhibits a great deal of variation mostly in theradiate sclerites from the surface cortex. The presence of 6-, 7-, and 8-radiates in this spe-cies is also equaled by P. arborea, though including quite different forms and size rangesin each species. In addition, P. johnsoni and P. arborea are the species with the largest geo-graphical and bathymetrical distributions within the group.

Distribution. Atlantic Ocean, Azores I. (Madeira), Brazil (off Rio de Janeiro), Floridaand Bahamas. 800–4152 m (Bayer 1964; Grasshoff, 1979).

Species comparisons. Grasshoff (1979) revised the type specimens of P. johnsoni andP. boschmai Bayer, including SEM analyses, and concluded that they belong to the samespecies, therefore P. boschmai is considered a junior synonymy of P. johnsoni. SinceGrasshoff’s (1979) work presented clear and straightforward information on species com-parisons no further attempts to describe the P. boschmai type specimen were undertaken.See also P. regalis Nutting and P. aotearoa sp. nov.


1014ZOOTAXA

FIGURE 10. Paragorgia johnsoni Gray (USNM 73767): A, stereo pair of a sclerite from the auto-zooid polyps (scale 10 µm); B, additional sclerites from the autozooid polyps (scales 20 µm); C–F,radiate sclerites (C–D are stereo pairs) from the colony surface (scales 10 µm); F, additional scler-ites from the colony surface and intermediates towards the medulla (scales 10 µm); G, scleritesfrom the colony medulla (scales 100 µm); H, sclerites from the subsurface/outer medulla (scales 20and 30 µm).


1014ZOOTAXAParagorgia splendens Thomson & Henderson, 1906

(Figs. 11–12)

Paragorgia splendens Thomson & Henderson 1906: 20; Bayer 1993: 4.Not Paragorgia sibogae Bayer 1993: 6.

Material examined. Holotype, BM(NH) 1933.3.13.38, British Museum Natural History,Indian Ocean off Sri Lanka (Investigator expedition st. 284, 7°55’N–81°47’E, 900 m).

FIGURE 11. Photograph of a fragment of the holotype of Paragorgia splendens Thomson &Henderson (courtesy of the British Museum-Natural History).


1014ZOOTAXA Diagnostic characters. Curved, slim terminal branches (up to 2 mm width). Surface

cortex (including calyx surface) containing mostly 8-rayed sclerites (radiate derived, likecapstans) up to 0.07 mm in length with rays formed by 4–6 blunt to pointed conical projec-tions (Fig. 12B–C).

FIGURE 12. Paragorgia splendens Thomson & Henderson, holotype: A, sclerites from the polyps(scale 10 µm); B–C, radiate sclerites from the colony surface (scales 10 µm) (B is a stereo pair); D,sclerites from the colony medulla (scales 100 µm).


1014ZOOTAXADescription. Fragile, gently bent, slim colonies with branches 1–2 mm in diameter at

the portions without autozooids (Fig. 11). Autozooids mostly isolated, occasional clustersof 3–4, along the branches within noticeable spherical calyces (Fig. 11). Autozooid polypapertures conical, semi-closed, projecting up to 2 mm. Colony easily breakable, most fea-tures closely resembling the original description drawing (Thomson & Henderson 1906).Color pale pink, medulla white. Medulla in the terminal branches visibly perforated by 2–

5 large circular internal canals (see also Bayer 1993). Polyps with blunt spindle-like scler-ites in the tentacles, up to 0.1 mm in length, with radially ornate belts of multiple lowcones mostly disorganized (Fig. 12A). Surface contains highly ornate, 8-radiate-derived,capstan-like sclerites (Fig. 12B–C), averaging 0.068 mm (±0.006 SD, n=10). Surfacesclerites 1.8 times longer than wider, averaging 0.038 mm in width (±0.003 SD, n=10).Medulla with long, slim, mostly straight, moderately ornate, spindles up to 0.7 mm inlength (Fig. 12D).

Morphological variation. The range of sizes is unknown for P. splendens because theoriginal material is very fragmented. The previous study of P. splendens did not includeSEM images of sclerites, but variation in both form and size among and within localities isoverall low (e.g., Bayer 1993: Fig. 2 vs. Fig. 12 this paper).

Distribution. Only known from the Indian Ocean off Sri Lanka (Investigator expedi-tion st. 284,7°55’N–81°47’E, 900 m, also station 333, see Bayer 1993).

Species comparisons. A record mentioned by Grasshoff (1979) from the Siboga sta-tion 95 (Sulu islands) was further described by Bayer (1993) as P. sibogae. However, it isimportant to note that P. sibogae Bayer was not included in this study because its descrip-tion was entirely based on a prepared sclerite slide (after Nutting 1911), without the assig-nation of a cataloged specimen. See also Paragorgia whero sp. nov.

Paragorgia regalis Nutting, 1912(Figs. 13–15)

Paragorgia regalis Nutting 1912: 100.Paragorgia dendroides Bayer 1956: 69 (sensu Bayer 1964: 526); Grasshoff 1979: 120.

Material examined. Holotype: USNM 30018, 34°15’N–138°E, 869–924 m, Honshuisland, Omae Zaki, 19 October 1906 (col. Albatross R/V), Saizuoka, Japan.

Other material: USNM 98789 (previously identified as P. dendroides by. F.M. Bayer),20° 46’ 57”N–157° 08’ 56”W, 1018 m, 20 September 1996 (Pisces DSR/V 5-301, LAD115), col. S. France & E. Bertson, Hawaii, USA; USNM 1014743 (previously identified asP. dendroides by. F.M. Bayer), 19 44 12N-158 17 43W, 452 m, Cross Seamount (RiverBasin), 5 February 2003 (col. R.B. Moffitt-OES 03-01 & E.H. Cave), USA.

Diagnostic characters. Autozooid polyps clustered in distinct and smooth globularnodules up to 2–3 mm high (Fig. 13). Surface cortex sclerites are small 7- and 8-radiates,


1014ZOOTAXA up to 0.08 mm including rays, with smooth surfaces. The small ones are greatly asymmet-

rical and integrating rays with distinctive small lumps (e.g., Figs. 14B–D, 15C–E).

FIGURE 13. Photograph of Paragorgia regalis Nutting (USNM 98789).

Description. Profusely branching colonies up to 300 mm in length with slender termi-nal branches down to 2–4 mm in diameter (Grasshoff 1979: Figs. 2–3). A few long, lateralbranches arise from the main branches and have a clavate appearance. Medulla perforatedby 3–4 large canals (Bayer 1956b: 69). Autozooid polyps on all sides of branches (Bayer1956b: 69) but with tendency towards one side (Fig. 13). Autozooids clustered in distinctglobular nodes up to 2–3 mm in height; siphonozooids much smaller (0.5 mm in height)but distributed throughout the surface (Bayer 1956b: 69). Salmon pink color. Autozooidpolyp tentacles contain ovals up to 0.1 mm in length, irregularly ornate with small conicaltubercles (Figs. 14A: 15A–B). Surface of the cortex, including autozooid apertures, withsmall 7- and 8-radiate sclerites up to 0.08 mm long but averaging 0.06–0.07 mm (±0.003


1014ZOOTAXASD, n=10, Holotype; ±0.005 SD, n=10, USNM 98789). Rays with smooth surfaces but

greatly asymmetrical in the smaller 7-radiate forms (Figs. 14B–D: 15C–E). Surface radi-ates about 1.6–1.7 times longer than wider, averaging 0.035–0.043 mm in width (±0.002–0.003 SD). The subsurface/outer medulla with larger, chiefly 8-radiates, up to 0.1 mm inlength (Bayer 1956b: 69). Medulla with ornate, coarsely warted spindles , oftentimesforked and ramified, up to 0.5 mm in length but frequently smaller (Figs. 14E; 15F).

FIGURE 14. Paragorgia regalis Nutting, holotype (USNM 30018): A, sclerites from the polyps(scales 10 µm); B–D, radiate sclerites from the colony surface (scales 10 µm) (B–C are stereopairs); E, sclerites from the colony medulla (scales 100 µm and 20 µm).


1014ZOOTAXA

FIGURE 15. Paragorgia regalis Nutting (USNM 98789): A–B, sclerites from the polyps (A is a

stereo pair) (scale 20 µm); C–E, radiate sclerites from the colony surface (scales 10 µm) (C–D are

stereo pairs); F, sclerites from the colony medulla (scales 100 µm).

Morphological variation. Surface sclerites, which include the diagnostic forms formost Paragorgia species, are quite variable in P. regalis. The sclerite drawings by Bayer(1956b: Fig. 1) are clearly the forms from the subsurface, which are usually large 8-radi-ates, and were also observed in the studied material (e.g., Fig. 14B–D). Since only two


1014ZOOTAXAspecimens were examined, a larger survey is needed to determine if Japanese specimens

(type locality) differ from Hawaiian populations (e.g., P. dendroides, Bayer 1993). Distribution. Pacific Ocean, Hawaii and Japan, 452–1018 m. Species comparisons. Bayer (1964: 526) was the first who suggested that P. dendroi-

des (Bayer 1956) should be a junior synonymy of P. regalis based on sclerite morphology.It is clear that sclerites from the type material of P. regalis (Japan) resemble those of theHawaiian specimens identified by F.M. Bayer as P. dendroides. Nonetheless, a detailedrevision of specimens throughout the Pacific is needed to clarify the synonymy of P. den-droides and/or the presence of a morphological cline across the Pacific. In comparisonwith other Paragorgia species, the 8-radiates from the subsurface/outer medulla of P.regalis resemble those from P. johnsoni and P. splendens, but the smaller forms in P.johnsoni are smoother without the irregular ornamentation observed in P. regalis and thecolonial features from P. splendens are quite different to those of P. regalis.

Paragorgia coralloides Bayer, 1993(Fig. 16)

Paragorgia coralloides Bayer 1993: 2.

Material examined. Holotype: USNM 50891, 01°54´00”S–127°36´00”E, 602 m, 03 Dec1909, (Albatross R/V), Obi Islands, Gomumu Island, Indonesia (Moluccas), Ceram Sea,

Diagnostic characters. Globular 8-radiate derived sclerites in the surface layer withsome radial ornaments subdivided in non-globular lobes (Fig. 16B–D).

Description. Colonies with slender branches down to 21 mm in diameter (Bayer1993). Autozooid polyps in clusters. Autozooid polyp tentacles contain spindles less than0.1 mm in length, irregularly ornate (Fig. 16A). Surface of the cortex, including autozooidaperture, with unusually globular and hypermorphic radiates (mostly 8-radiate derived) upto 0.055 mm averaging 0.0435 mm in length (±0.003 SD, n=10) (Fig. 16B–D). Surfaceradiates about 1.638 times longer than wider, averaging 0.029 mm in width (±0.002 SD,n=10). Despite radiates have some globular and larger rays, one side of radiates remainswith smaller multilobular ornaments. Medulla with ornate spindles, rather short, up to 0.15mm in length (Fig. 16E).

Morphological variation. The number of radial ornaments of the sclerites of the sur-face layer, a diagnostic character, can be highly variable in number but most radiatespresent such a character.

Distribution. Pacific Ocean: Palau Gomumu (Obi Major) (Bayer 1993). Species comparisons. Bayer (1993) suggested that P. coralloides radiates are indistin-

guishable from those from Corallium spp. Although there is a great resemblance, P. coral-loides do not form asymmetrical opera-glass sclerites (e.g., Bayer 1956: Fig. 7), wheretypically lateral rays are enlarged. P. coralloides usually have the proximal and distal rays


1014ZOOTAXA enlarged. Nonetheless, the tentacle sclerites somewhat resemble those from Corallium

spp., which brings up a possible phylogenetic connection between Corallium and Para-gorgia. See also P. tapachtli sp. nov.

FIGURE 16. Paragorgia coralloides Bayer (USNM 50891): A, sclerites from the polyps (scales

10 and 20 µm); B–D, radiate sclerites from the colony surface (scales 10 µm) (C–D are stereo

pairs); E, sclerites from the colony medulla (scales 20 µm).


1014ZOOTAXAParagorgia alisonae sp. nov.

(Figs. 17–21)

Material examined. Holotype: NIWA 3312, H-842, J213, Z9596, 48° 01’S–166° 05’E,“Otara hill”, 980 m, 27 November 1998 (col. J. Wills, FV Amatal Explorer 1171/25, bot-tom trawl).

Paratypes : NIWA 3313, P-1395, J224 and NIWA 3314, P-1396, J244 same as holo-type locality; NIWA 3315, P-1397, J211, Z9595, 48° 01’S–166° 06’E, “Otara hill”, 940–1180 m, 27 November 1998 (col. J. Wills, FV Amatal Explorer, 1171/24, bottom trawl);NIWA 3316, P-1398, J248, Z8981, 44° 57.68’–56.75’S – 174° 11.23’–09.56’E, south-westChatham Rise, 1041–1052 m, 5 December 1997 (RV Tangaroa 9713/37, bottom trawl);NIWA 3317, P-1399, J250, Z9583, 48° 02.1' S–166° 06.1' E, “Doghill seamount”, 935m,25 Nov 1998 (col. J. Wills, FV Amatal Explorer 1171/12, bottom trawl).

Diagnostic characters. Surface of the cortex (including calyx surface) contains highlyornate sclerites (radiate derived, like capstans), oval shaped, up to 0.09 mm in length withmultiple and particular blunt lobes on the rays (Fig. 18D–F, 19D–F).

Description. Large colonies, holotype and paratypes composed of multiple fragments< 200 mm in maximum length (Fig. 17A–B). Some colonies (paratypes) up to a meter inheight and 43 mm in diameter (e.g., Fig. 17C–D). Robust branches down to 3–5 mm at theslimmer parts (Fig. 17B). Colonies with simple lateral branching; every major branch car-rying multiple short lateral branches, as observed in most octocorals (e.g., Sánchez et al.2004), tending upward and in one-plane, with branch tips usually ending in a bulb. Colonysurface in high relief, though some colonies (paratypes) have autozooids toward one sideof colony. Numerous conical, semi-closed, autozooid polyp apertures up to 4 mm in diam-eter with the tendency to irregularly aggregate as nodules (Fig. 17). Bright red color,white/red mixed medulla, extended polyps pale yellow. Medulla in the terminal brancheswith 3–8 major canals plus numerous smaller canals; subsurface also with small canals.Autozooid polyp tentacles with ovals, similar to other Paragorgia spp., up to 0.13 mmlong, regularly but not profusely ornate with smooth conical rays (Figs. 18A–C, 19A–C)and sometimes shorter and profusely ornate (e.g., Fig. 18B). Surface sclerites oval, highlyornate, radiate-derived, capstan-like sclerites (Fig. 18D–E, 19D–E), averaging 0.078 mmin length (±0.005 SD, n=10, NIWA 3312; ±0.006 SD, n=10, NIWA 3315). Oval surfacesclerites 1.5 times longer than wide, averaging 0.05 mm in width (±0.002 SD, n = 10,NIWA 3312 = 3315). Subsurface with intermediate forms to longer spindle-like sclerites(Figs. 18D, 19D). Medulla with long and slim, slightly bent but mostly straight, moder-ately ornate, spindles up to 0.4 mm in length (Figs. 18H, 19G).

Morphological variation. The two specimens used for the description, NIWA 3312and 3315, are nearly identical in all characters with overall low variation probably becausethey came from the same locality. However, specimens NIWA 3316–17 from differentlocations comprised larger and thicker specimens, almost resembling P. arborea colonies,Fig. (17C–D), and with autozooid mostly located towards one side. The sclerites, although


1014ZOOTAXA not identical to those from the specimens from the type locality, fall within the overall

forms and characters for P. alisonae sp. nov. (Figs. 20–21).

FIGURE 17. Paragorgia alisonae sp. nov.: A–B, holotype (NIWA 3312); C–D, paratypes (NIWA3316–3317 respectively). Calliper opened at 2 cm.


1014ZOOTAXA

FIGURE 18. Paragorgia alisonae sp. nov., paratype (NIWA 3315): A–B, stereo pairs of the scler-ites from the polyps (scale 10 µm); C, additional sclerites (3) from the polyps (scales 20 µm); D–F,radiate sclerites from the colony surface (scales 10 µm) (E and F are stereo pairs); G, sclerites fromthe colony medulla (scales 100 µm).


1014ZOOTAXA

FIGURE 19. Paragorgia alisonae sp. nov., holotype (NIWA 3312): A–B, stereo pairs of the scler-ites from the polyps (scales 20 and 10 µm); C, additional sclerites (2) from the polyps (scales 10µm); D–F, radiate sclerites from the colony surface (scales 10 µm) (E and F are stereo pairs); G,sclerites from the colony medulla (scales 100 µm).


1014ZOOTAXA

FIGURE 20. Paragorgia alisonae sp. nov., paratype (NIWA 3316): A–B, stereo pairs of the scler-ites from the polyps (scales 10 and 20 µm); C–D, additional sclerites from the polyps (scales 20µm); E–G, radiate sclerites from the colony surface (scales 10 µm) (F and G are stereo pairs); H,sclerites from the colony medulla (scales 100 µm).

Distribution. P. alisonae is known from the type locality at 980 m depth, from TheSnares, south of Stewart Island, southwest of the Chatham Rise, and Dog Hill seamount alllocations in the New Zealand EEZ around 935–1180 m (Fig. 2).

Species comparisons. P. alisonae is related to P. splendens in many aspects but theirdifferences are also straightforward. These two species present similar surface sclerites as


1014ZOOTAXA well as pale yellow extended autozooid polyps, projecting from similar conical structures.

However, P. alisonae has more robust terminal branches than P. splendens, sometimes asthick as P. arborea. Surface sclerites, a diagnostic character, are near perfectly oval in P.alisonae (1.5 times longer than wider) whereas in P. splendens these are enlarged (1.7times) with less ray ornaments. It is important to mention that under optical microscopethe surface sclerites from these two species are nearly identical and SEM is highly recom-mended to further tell them apart.

Etymology. The species name is dedicated to my wife Alison R. Acosta-de-Sánchez.

FIGURE 21. Paragorgia alisonae sp. nov., paratype (NIWA 3317): A, stereo pair of a sclerite from

the polyps (scale 10 µm); B, additional sclerites from the polyps (scales 10 µm); C–E, radiate scler-

ites from the colony surface (scales 10 µm) (C and D are stereo pairs); F, sclerites from the colony

medulla (scales 100 µm).


1014ZOOTAXAParagorgia kaupeka sp. nov.

(Figs. 22–25)

Material examined.Holotype: NIWA 3318, H-843, J204, Z10061, 37° 02.8’S–176°29.8’E, “No name knoll”, 949 m, 27 March 2000 (col. D. Wrightson, FV Ocean Fresh1337/12, bottom trawl).

Paratypes: NIWA 3319, P-1400, J205, Z10061-dry, same as holotype; NIWA 3320, P-1401, J215, X152, 36° 09.74’S–176° 48.38’E, “Colville knolls”, 820–940 m, 28 Novem-ber 1989 (RV Rapuhia 2032, rock dredge); NIWA 3321, P-1402, J207, Z9229, 36° 53.8’S–

177° 22.3’E, “Otara Knoll” 787 m, 15 August 1998 (col. J. Wills, FV Margret Phillipa1124/65, bottom trawl).

Diagnostic characters. Surface cortex, including autozooid aperture, with small,rounded, radiate sclerites less than 0.05 mm in length, averaging 0.04 mm, 1.65–1.7 timeslonger than wide, losing the radiate symmetrical form by enlarging most radial ornaments:surface irregular and uneven (Figs. 24C–D, 25D–F).

FIGURE 22. Paragorgia kaupeka sp. nov., holotype (NIWA 3318): A, entire specimen; B, side ofbranches with autozooids; C, opposed side without autozooids; calliper open at 20 mm.


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FIGURE 23. Paragorgia kaupeka sp. nov., paratype (NIWA 3319): A, entire specimen; B, side of

branches without autozooids; C, opposed side with autozooids; calliper open at 20 mm.

Description. Branching colonies, pseudo-dichotomous, uniplanar, up to 40 cm inheight (holotype 30 cm: Fig. 22). Slim terminal branches 4–6 mm in diameter (Figs. 22–

23). Autozooids usually clustered in groups of 2–7 polyps, towards one face of the colony,as alternately distributed hemispheric nodules on the branches (Figs. 22–23). Surface cor-tex with numerous tiny granular siphonozooid apertures present on both sides of the col-ony. Colonies pale red in alcohol and pink when dried, medulla predominately white.Autozooid polyps often retracted but occasionally extended in preserved samples exhibit-ing pale yellow tentacles. Medulla in the terminal branches perforated by 4–6 main canalsand usually including red sclerites. Subsurface exhibiting 18–22 clearly discernable canalsaround the medulla. Tentacles with common Paragorgia sclerites: ornate blunt ovals up to0.1 mm in length, usually with cone-like prominences and a smooth waist (Figs. 24A–B,25A–B), and with occasional twinned forms producing crosses (Fig. 25C). Surface of thecortex, including autozooid aperture, with small, radiate sclerites less than 0.05 mm inlength averaging 0.04 mm (±0.0028 SD, n=10, NIWA 3319; ±0.0038 SD, n=10, NIWA3318) (Figs. 24C–D; 25D–F). Surface radiates between 1.65–1.7 times longer than wide,averaging 0.02 mm in width (±0.0028 SD, n=10, NIWA 3319; ±0.0035 SD, n=10, NIWA3318). They show a loss of the radial symmetry by an asymmetrical enlargement of most


1014ZOOTAXAradial ornaments (Figs. 24C–D; 25D–F); the uneven and irregular form and contours of the

enlarged radial ornaments include considerably larger amounts of calcite than shorter rayornaments. Subsurface sclerites include the same radiates as the surface with the occa-sional presence of larger forms with similar shapes and short, smooth but ornate, spindlesup to 0.25 mm. Medulla with long, slim, mostly straight but occasionally bent, ornate,spindles less than 0.4 mm in length usually up to 0.35 mm (Figs. 24E, 25G).

FIGURE 24. Paragorgia kaupeka sp. nov., holotype (NIWA 3318): A, stereo pair of a sclerite fromthe polyps (scale 10 µm); B, additional sclerites from the polyps (scales 10 µm); C–D, radiate scler-ites (C is a stereo pair) from the colony surface (scales 10 µm); E, sclerites from the colony medulla(scales 100 µm).


1014ZOOTAXA

FIGURE 25. Paragorgia kaupeka sp. nov., paratype (NIWA 3321): A–C, stereo pairs of the scler-ites from the polyps (scale 10 µm); D–F, radiate sclerites (stereo pairs) from the colony surface(scales 10 µm); G, sclerites from the colony medulla (scales 100 µm).


1014ZOOTAXAMorphological variation. Owing to the irregular ornamentation of the diagnostic cor-

tex surface sclerites, there was quite a lot of individuality in the form of the radiates amongand within specimens (e.g., Figs. 24C–D, 25D–F).

Distribution. North of the Bay of Plenty, New Zealand, between 36°09’–37°2.8’S and176°29.8’–177°22.3’E, in an area rich in seamounts and underwater volcanoes (Fig. 2),747–949 m depth.

Species comparisons. P. kaupeka sp. nov. colonies resemble those of P. johnsoni(Grasshoff, 1979), particularly in the branching pattern and the autozooid clusters arrangedtowards one side. However, Caribbean samples of P. johnsoni (=P. boschmai) in terminalbranches are definitely thinner (up to 3 mm in diameter: Bayer, 1964) than P. kaupeka (4–

6 mm). P. johnsoni has most of the radiates regularly ornate and smoother (Grasshoff,1979: Fig. 5) than P. kaupeka.

Etymology. The species is named after the Maori word kaupeka, meaning “branch” ofa tree, alluding to the branching nature of these octocorals.

Paragorgia maunga sp. nov.(Figs. 26–27)

Material examined.Holotype: NIWA 3322, H-844, J210, Z10989, 33° 53.2’S–167°56.3’E, “Wanganella Bank” 1082 m, 22 January 2002 (col. S. Smith, F/V Waipori, bottomtrawl).

Paratypes: NIWA 3323, P-1403, J223, Z9779, 34° 07.1–07.55’S–174° 53.7–54.0’E,“Seamount #441” 808–1121 m, 25 June 1999 (col. C. Blincoe, FV Seamount Explorer1236/26, bottom trawl); NIWA 3324, P-1404, J67, 35° 43.71' S, 178° 29.98' E, “RumbleIII seamount”, 545–671m, 3 November 2000 (RV Kaharoa 0011/28, epibenthic sled).

Diagnostic characters. Delicate, soft, red cortex. Medulla perforated by only 2–3small axial canals, densely surrounded by red sclerites; numerous even smaller canalsthroughout. Surface sclerites with an asymmetrical enlargement of most radial ornamentsgenerating smoothly lobulated sub-ornaments in all rays.

Description. Colonies with a delicate, thin, soft, red cortex; all examined material par-tially eroded exhibiting the white medulla on surface. The holotype is a fragmented colony210 mm in length (Fig. 26); the paratypes are broken fragments less than 120 mm long.Branching possibly pseudo-dichotomous in one plane (no complete specimens available),with autozooid polyps towards one face. Autozooids forming irregular clumps of 3–5 pol-yps (Fig. 26B). Slimmer parts of branches down to 2–3 mm width. Medulla perforated byonly 2–3 main small canals, tending central, and numerous even smaller canals in both themedulla and subsurface/outer medulla. Center of the medulla, around the main canals,with red sclerites, otherwise colorless with occasional red ones. Autozooid tentaclesexhibiting the common red Paragorgia sclerites: ornate ovals up to 0.1 mm in length withconical ornaments throughout (Fig. 27A–C, J). Surface of the cortex, including autozooid


1014ZOOTAXA aperture, with small, radiate sclerites up to 0.08 mm in length averaging 0.06–0.07 mm

(±0.006 SD, n=10, NIWA 3322=3324) (Fig. 27D–H, K). Surface radiates between 1.7–1.8times longer than wide, averaging 0.04 mm in width (±0.004 SD, n=10, NIWA3322=3324). Asymmetrical enlargement of most radial ornaments generating smoothlylobulated sub-ornaments in all rays (Fig. 27D–H, K). Medulla with long, straight, orna-mented (occasional large, irregular projections) spindles up to 0.35 mm in length (Fig.27I).

FIGURE 26. Paragorgia maunga sp. nov., holotype (NIWA 3322): A, whole colony; B, branchdetail (calliper open at 20 mm).

Morphological variation. There were some small (<0.01 mm) but significant differ-ences in the size of surface sclerites of two specimens (t=5.5, df=19, P<0.001, only 10sclerites measured in each specimen) although no variance differences were found amongthe specimens (F=0.8, df=9, P>0.4). Since there is an expected variation in the surfacesclerites, i.e., progressively larger towards the subsurface, and the examined specimenswere highly eroded during the collection, the differences among specimens could be due todifferential sampling in relation to the surface-subsurface. Nonetheless, all examined spec-imens clearly exhibit the species diagnostic characteristics (e.g., Fig. 27A–I vs. J–K).

Distribution. The three specimens examined span a large area of the EEZ of NewZealand from the West Norfolk ridge to two seamounts North off the Bay of Plenty (Fig.1).


1014ZOOTAXA

FIGURE 27. Paragorgia maunga sp. nov., A–I, holotype (NIWA 3322): A, stereo pair of a scleritefrom the polyps (scale 10 µm); B–C, additional sclerites from the polyps (scales 10 and 20 µm); D–H, radiate sclerites (D–G stereo pairs) from the colony surface (scales 10 µm); I, sclerites from thecolony medulla (scales 100 µm). J–K, Paratype (NIWA 3324): J, sclerite from the polyps (scale 30µm); K, radiate sclerites from the colony surface (scales 10 µm).


1014ZOOTAXA Species comparisons. Externally and in the shapes of the surface sclerites, P. maunga

sp. nov. resembles P. kaupeka, and a close phylogenetic relationship is expected amongthese two species. However, it is rather easy to tell them apart by comparing their diagnos-tic characters. The form of the surface radiate sclerites can seem nearly identical under thelight microscope but there is nearly 0.03 mm difference in the mean length of the two spe-cies’ sclerites, which can be determined using a micrometer and compound microscope.Under SEM it is also evident that the enlargement of the radial ornamentation in the sur-face sclerites of P. kaupeka is greatly irregular, generating a fairly rough surface, whereasin P. maunga it is smooth and lobate.

Etymology. The word maunga means “mount” in Maori, alluding to the collection ofspecimens from seamounts.

Paragorgia aotearoa sp. nov.(Figs. 28–29)

Material examined. Holotype: NIWA 3325, H-845, J206, 42°50.26’–42°49.99’S,176°54.96’–176° 55.17’W, “Mt Muck seamount” 700–900 m, August 1996, (RV Tangaroa9609/40, bottom trawl, col. D. Tracey).

Diagnostic characters. Medulla with ornate but smooth and blunt spindles usuallyless than 0.3 mm in length. Surface of the cortex, including autozooid aperture, with com-paratively (among Paragorgia spp.) large 8-radiate sclerites up to 0.09 mm.

Description. Profusely branching colonies up to 230 mm in length (holotype) with ter-minal branches down to 3–4 mm in diameter (Fig. 28A). Numerous short, lateral branchesarise from the main branches with an irregular but clavate appearance (Fig. 28A). Autozo-oid (and siphonozooid) polyps located towards one side of the colony (Fig. 28B) whereasthe other side has only siphonozooids (Fig. 28C). There is a regular trend for the autozoo-ids to form clumps, but many autozooids are several millimeters apart from each other.Cortex red and thick (~>0.5 mm) containing siphonozooids fairly uniformly-distributed~0.5 mm apart. Medulla perforated by 3–4 main axial canals at terminal branches, usuallysurrounded by red sclerites. Autozooid polyp tentacles with blunt ovals, slightly thickerthan in most Paragorgia spp., up to 0.1 mm in length and 0.5 mm width, regularly andprofusely ornate with conical tubercles (Fig. 29A–C). Surface of the cortex, includingautozooid aperture, with comparatively (among Paragorgia spp.) large 7- to 8-radiatesclerites up to 0.09 mm averaging 0.076 mm in length (±0.008 SD, n=10) (Fig. 29D–F).Rays nearly symmetrical with smooth surfaces but occasional lobulated (Fig. 29D–F). Sur-face radiates between 1.65 times longer than wider, averaging 0.04 mm in width (±0.004SD, n=10). Medulla with ornate (red and colorless) but smooth and blunt spindles up to0.33 mm in length averaging 0.28 mm (±0.03 SD, n=10) (Fig. 29G). Holotype with a com-mensal white anemone.


1014ZOOTAXA

FIGURE 28. Paragorgia aotearoa sp. nov., holotype (NIWA 3325): A, whole colony; B, branchdetail apolypar side; C, polypar (autozooid) side (calliper open at 20 mm).

Morphological variation. Unfortunately, it is not possible to make specimen compar-isons since only one specimen of P. aotearoa sp. nov. was available. Overall, intracolonialvariation of sclerites and other features is very low, with rather low variation in surfaceradiates: a character where most of the variation in form is present in Paragorgia spp.

Distribution. Only known for the type location, Chatham Rise between the NewZealand’s North Island and the Chatham islands.

Species comparisons. Some features of P. aotearoa closely resemble P. johnsoni Grayand P. regalis Nutting. For instance, the sclerites from the cortex surface are chiefly 8-radi-ates with smooth but slightly lobate radial ornamentation (Figs. 17D–F; 18B) (see alsoBayer, 1956: Fig. 1). However, the diagnostic character of P. aotearoa, short medullarsclerites up to 0.3 mm, are surpassed by P. regalis with spindles up to 0.4 mm (Fig. 27C).In addition, P. regalis colonies are salmon pink (Bayer, 1956; pers. obs.) with largerbranches and polyps (autozooids/siphonozooid) (e.g., Fig. 13). The surface sclerites of P.aotearoa are very similar in shape to the surface sclerites of P. johnsoni (Fig. 10C–F).However, P. aotearoa sclerites are clearly larger than those of P. johnsoni (0.08 vs 0.05mm in average respectively); in fact, P. aotearoa has the largest surface sclerites in all spe-cies of Paragorgia. Autozooid polyp sclerites of P. johnsoni also differ from those of P.


1014ZOOTAXA aotearoa in the irregular ornamentation. Nonetheless, the phylogenetic affinity between P.

regalis, P. johnsoni, and P. aotearoa is clear.

FIGURE 29. Paragorgia aotearoa sp. nov., holotype (NIWA 3325): A–B, stereo pairs of the scler-ites from the polyps (scale 10 µm); C, additional sclerites from the polyps (scales 20 and 10 µm);D–F, radiate sclerites from the colony surface (scales 10 µm) (E and F are stereo pairs); G, scleritesfrom the colony medulla (scales 20 µm).


1014ZOOTAXAEtymology. The word Aotearoa means in Maori, “the land of the long white cloud” as

the first Maori settlers called what is today known as New Zealand.

Paragorgia wahine sp. nov.(Figs. 30–31)

Material examined. Holotype: NIWA 3326, H-846, J247, 42°47.47' S–179°59.33' W,“Diabolical seamount”, 890–1000m, 17 April 2001 (RV Tangaroa 0104/113, epibenthicsled).

Diagnostic characters. Slim branches up to 2 mm in diameter at the portions withoutautozoids. Surface sclerites highly ornate, 8-radiate, capstan-like sclerites (Fig. 31), aver-aging 0.07 mm and 1.7 times longer than wide, averaging 0.04 mm in width. Medulla withlong, slim, highly ornate, spindles up to 0.3 mm.

Description. The holotype is a fragile and slim single branch up to 2 mm in diameterat the portions without autozoids (Fig. 30). Autozoids with gregarious tendency, with clus-ters up to 7 mm in diameter, but occasionally isolated on the other portions of the branch(Fig. 30). Autozooid polyps fully retracted. Autozooid polyp apertures low. Colony easilybreakable, 50 mm in length (Fig. 30). Color variable between pink and red. Medulla in theterminal portion of the branch perforated by 5–7 small, circular, internal canals. Polypswith blunt, long spindles in tentacles, up to 0.1 mm in length, with radial ornate belts ofmultiple acute cones (Fig. 31A–C). Surface sclerites highly ornate, 8-radiate-derived, cap-stan-like sclerites (Fig. 31D–H), averaging 0.07 mm (±0.004 SD, n=10). Surface sclerites1.7 times longer than wide, averaging 0.04 mm in width (±0.002 SD, n=10). Subsurfacewith spindle-like sclerites of intermediate form. Medulla with long, slim, highly ornate,spindles up to 0.3 mm in length (Fig. 31I).

Figure 30. Paragorgia wahine sp. nov., holotype (NIWA 3326): colony fragment and detail (calli-per open at 20 mm).


1014ZOOTAXA Morphological variation. Unfortunately, only one specimen was available for this

species. It was possible to see a great deal of variability in the ornamentation of the surfaceand medulla sclerites.

FIGURE 31. Paragorgia wahine sp. nov., holotype (NIWA 3326): A–B, stereo pairs of the scler-

ites from the polyps (scale 10 µm); C, additional sclerites from the polyps (scale 20 µm); D–H,

radiate sclerites from the colony surface (scales 10 µm) (D to G are stereo pairs); I, sclerites from

the colony medulla (scales 100 µm).


1014ZOOTAXADistribution. Known only from “Diabolical Seamount”, east of New Zealand.

Species comparisons. All the sclerites from P. wahine sp. nov. are similar to thosefrom P. splendens. However, P. splendens surface sclerites are overall larger than P.wahine. Surface sclerites average 0.089 mm in P. splendens compared to 0.071 in P.wahine. Likewise, medullar sclerites are up to 0.5 mm in P. splendens and usually less than0.3 mm in P. wahine. More obvious differences between the two species are in the exter-nal appearance, where P. wahine is certainly slimmer and smoother than P. splendens.

Etymology. Wahine, “bride”, “wife” or “woman” in Maori, was an inter-island ferry inNew Zealand that sunk on 10 April, 1968. The species is named in remembrance of all thevictims of the disaster.

Paragorgia whero sp. nov.(Figs. 32–33)

Material examined.Holotype: NIWA 3436, H-875, J214, Z9583, 48 °02.1' S, 166 ° 06.1'E, “Doghill seamount”, 935m, 25 Nov 1998 (col. J. Wills, FV Amatal Explorer 1171/12,bottom trawl).

Paratype: NIWA 3439, P-1427 (J74), 36° 8.84' S, 178° 12.24' E, “Rumble V sea-mount”, 772–951 m, 23 May 2001 (RV Tangaroa 0107/225, epibenthic sled).

Diagnostic characters. Cortex surface (including calyx surface) containing mostly 8-rayed sclerites (radiate derived, like capstans) up to 0.09 mm in length with rays formed by4–6 blunt to pointed, conical projections (Fig. 33D–E).

FIGURE 32. Paragorgia whero sp. nov.: A–B, holotype (NIWA 3436); C, Paratype (NIWA 3439).

Description. Fragile, slim colonies with branches 3–4 mm in diameter at the portionswithout autozooids (Fig. 32). Autozooids with gregarious tendency in the branches withclusters up to 7 mm in diameter, but also isolated on slimmer portions of the branches (Fig.32). Autozooid polyps extended in alcohol-preserved specimens; sweeper tentacles, paleyellow. Autozooid polyp apertures conical, semi-closed, projecting up to 2 mm from thebranch. Colony easily breakable, larger fragments 63 mm in length (holotype: Fig. 32A).


1014ZOOTAXA

FIGURE 33. Paragorgia whero sp. nov., holotype (NIWA 3436): A–B, stereo pairs of the scleritesfrom the polyps (scale 10 µm); C, additional sclerites from the polyps (scales 20 µm); D–F, radiatesclerites from the colony surface (scales 10 µm) (E and F are stereo pairs); G, sclerites from the col-ony medulla (scales 20 µm).


1014ZOOTAXAColor bright red (NIWA 3436) to pink (NIWA 3439), with a white medulla. Medulla at the

terminal branches visibly perforated by 2–5 large circular internal canals. Polyps withblunt spindles in the tentacles, up to 0.1 mm in length, with radially ornate belts of multi-ple acute cones (Fig. 33A–C). Surface sclerites highly ornate, 8-radiate-derived, capstan-like sclerites (Fig. 33D–E), averaging 0.089 mm (±0.005 SD, n=10, NIWA 3436; ±0.008SD, n=10, NIWA 3439). Surface sclerites 1.65–1.74 times longer than wide, averaging0.05 mm in width (±0.004 SD, n=10, NIWA 3436; ±0.006 SD, n=10, NIWA 3439). Sub-surface with forms intermediate to longer spindle-like sclerites (Fig. 33D). Medulla withlong, slim, mostly straight, moderately ornate, spindles up to 0.5 mm in length (Fig. 33G).

Morphological variation. Owing to the small and fragmented nature of the studiedmaterial the full range of sizes for P. whero sp. nov. is unknown. In any case, the branchesare so fragile it is unlikely that the species attains a large size.

Distribution. Doghill and Rumble V seamounts, 772–951 m. Species comparisons. P. whero has more slender branches (3–4 mm) and autozooid

nodules compared to P. arborea (10–30 mm) but thicker branches than P. splendens. Thesclerites from the surface cortex (other layers are less variable across species) exhibitmostly 8-radiates of larger size, which was among the characters chosen by Bayer (1993)as diagnostic for P. splendens, suggesting relatedness between the two species. Bayer(1993) also reported some hypertrophy (e.g., Opera glass type) but that was not observedin P. whero.

Etymology. The word “whero” means red in Maori, one characteristic of this species.

Paragorgia tapachtli sp. nov.(Figs. 34–35)

Material examined. Holotype: USNM 98786, 12° 44’ N–102° 36’ W, 1950 m, (Seamount6, Alvin DSR/V, P. Hickey), 29 October 1995, Mexico.

Diagnostic characters. Autozooid polyp tentacles with irregularly ornate spindles androds, slightly different than in most Paragorgia spp., less than 0.1 mm in length (Fig. 35A–

B). Cortex surface, including autozooid aperture, with unusually globular and hypermor-phic radiate (mostly 6-radiate derived) sclerites up to 0.047 mm.

Description. Colony branching in one plane, slender branches, slightly curved, downto 2 mm in diameter (Fig. 34). Autozooid polyps distributed uniformly with a notably con-ical aperture (Fig. 34). Colony bright red. Medulla perforated by 1–2 main axial canals inthe terminal branches. Autozooid polyp tentacles with irregularly ornate spindles rods,slightly different to most Paragorgia spp., less than 0.1 mm in length (Fig. 35A–B). Sur-face of the cortex, including autozooid apertures, with unusually globular and smooth radi-ate (mostly 6-radiate derived) sclerites up to 0.047 mm averaging 0.043 mm in length(±0.003 SD, n=10) (Fig. 35C–F). Surface radiates about 1.63 times longer than wide, aver-aging 0.02 mm in width (±0.001 SD, n=10). Medulla with ornate spindles up to 0.3 mm inlength (Fig. 35G).


1014ZOOTAXA Morphological variation. Both diagnostic characters, autozooid tentacle and surface

sclerites, exhibit a great deal of variation in P. tapachtli sp. nov. The globular radiates ofthe surface present particular individualized forms but are always smooth with little radialornamentation.

Distribution . Off Mexico, eastern Pacific Ocean, 1950 m.Species comparisons. No other Paragorgia species has such highly derived charac-

ters as P. tapachtli, though they are similar in P. coralloides but not as exaggeratedly glob-ular and rounded.

Etymology. Tapachtli is the word used for ‘coral’ in Nahuatl tongue, an indigenouslanguage from Mexico, the country close to the place where this species was discovered.

FIGURE 34. Paragorgia tapachtli sp. nov., holotype (USNM 98786) (colony is 20 cm height and3.5 mm of basal branch diameter).


1014ZOOTAXA

FIGURE 35. Paragorgia tapachtli sp. nov., holotype (USNM 98786): A–B, sclerites from the pol-yps (scales 20 µm); C–F, radiate sclerites from the colony surface (scales 10 µm) (C–D–E are stereopairs); G, sclerites from the colony medulla (scales 100 µm).

Paragorgia yutlinux sp. nov.(Figs. 36–38)

Material examined.Holotype: USNM 1073480, BC24, 50°13’54”N–128°35’00”W, offBritish Columbia and Vancouver island, 846–861 m, (E/W Ricker, col. J. Boutillier), 12April 2003, Canada.


1014ZOOTAXA Paratype: USNM 57217, 47°35’48”N–125°08’42”W, 503 m, ‘John Cobb seamount’,

(John N. Cobb R/V, AEC Project), 16 March 1962, Washington, USA.Diagnostic characters. Colonies with a white cortex and pink/purple autozooid aper-

tures. Numerous conical, semi-closed, autozooid polyp apertures uniformly/randomly dis-tributed throughout the branches. Surface sclerites are mostly 6-radiates with long,lobulated, and smooth rays (Fig. 37D–G, 38D–I).

FIGURE 36. Paragorgia yutlinux sp. nov.: A, holotype (USNM 1073480) (colony fragment is 7.5cm long with a branch diameter of 6.8 mm); B, Paratype (USNM 57217).

Description. Robust colonies, with white cortex and pink to purple autozooid aper-tures. Numerous conical, low, semi-closed, autozooid polyp apertures uniformly/randomlydistributed throughout the branches (Fig. 36). Tiny siphonozooid apertures tightly closed,not observable with the naked eye. Medulla in terminal branches, with 3–5 major canals.


1014ZOOTAXAAutozooid polyp tentacles with blunt ovals, similar to other Paragorgia spp., up to 0.13

mm long, regularly ornate with small conical tubercles (Figs. 37–38A–C). Surface scler-ites mostly 6-radiates with long lobulated and smooth rays (Fig. 37D–G, 38D–I), averaging0.046–0.054 mm in length (±0.008 SD, n=10, USNM 57217; ±0.003 SD, n=10, USNM1073480). Surface sclerites between 1.36–1.54 times longer than wide, averaging 0.03 mmin width (±0.005 SD, n=10, USNM 57217; ±0.002 SD, n=10, USNM 1073480). Subsur-face with forms intermediate to longer spindle-like sclerites (e.g. Fig. 38I). Medulla withslim, straight, ornate, spindles usually less than 0.25 mm in length (Figs. 37H, 38J).

FIGURE 37. Paragorgia yutlinux sp. nov., holotype (USNM 1073480): A–C, sclerites from thepolyps (scales 10 µm) (A–B are stereo pairs); D–G, typical surface sclerites (scales 10 µm) (D–Fare stereo pairs); H, sclerites from the medulla (scales 20 µm).


1014ZOOTAXA Morphological variation. The sizes of the diagnostic sclerites, 6-radiates, showed a

small but significant difference between the two examined specimens (t=2.95, df=18,P<0.01), although no notable differences in form occur (e.g., Figs. 37–38). Because of thesmall number of samples is not possible to tell if the variation is due to a gradient or not.

Distribution. Eastern Pacific Ocean: off British Columbia (Canada) and Washington(USA), 503–1000 m.

FIGURE 38. Paragorgia yutlinux sp. nov., paratype (USNM 57217): A–C, typical polyp sclerites(scales 10 µm) (A–B are stereo pairs); D–H, radiates from the surface (scales 10 µm); I, intermedi-ate form between the surface and internal medulla (scale 20 µm); J, sclerites from the medulla(scales 100 µm).


1014ZOOTAXASpecies comparisons. Interestingly, externally this species is similar to species of

Sibogagorgia due to the absence of aggregations of autozooids, although it is clear that P.yutlinux sp. nov. is not from that genus. The other Paragorgia species with mostly 6-radi-ates is P. arborea, but they are considerably smaller than P. yutlinux with different orna-mentation as well. The radial ornamentation of P. yutlinux sp. nov. 6-radiates can resemblethe 7- and 8-radiates from P. alisonae but their morphologies are definitely divergent andunique in each case.

Etymology. This species is named in honour of the vanished Kwakwaka'wakw tribe,Yutlinux, from Cox and Lanz Islands, British Columbia, Canada.

Paragorgia stephencairnsi sp. nov.(Figs. 39–41)

Material Examined.Holotype: USNM 57982, 350 m, col. N. McDaniel, exact localityunknown, off British Columbia, Canada.

Paratype: USNM 94437, 490 m, Alvin DSR/V (Dive 2296), 16 October 1990,32°26’00”N–127°47’36”W, California, USA.

Diagnostic characters. Colonies with flattened branches. Surface sclerites mostly 7-radiates (occasionally 8-radiates) with long (> 0.01 mm) lobulated, smooth rays (Fig.40D–G, 41B–D), averaging 0.065 mm in length. Medullar sclerites highly ornate, forked,irregular, spindles usually less than 0.3 mm in length (e.g., Fig. 40H).

FIGURE 39. Paragorgia stephencairnsi sp. nov., holotype (USNM 57982) (the colony fragment is

6.5 cm in height).


1014ZOOTAXA

FIGURE 40. Paragorgia stephencairnsi sp. nov., holotype (USNM 57982): A–B, typical polypsclerites (scales 10 µm) (A is a stereo pair); C–G, radiates from the surface (scales 10 µm) (C–E–Gare stereo pairs); H–I, sclerites from the medulla (scales 100 µm) (H is a stereo pair) (scale 30 µm).


1014ZOOTAXA

FIGURE 41. Paragorgia stephencairnsi sp. nov., paratype (USNM 94437): A, typical polyp scler-

ites (scales 10 µm); B–D, radiates from the surface (scales 10 µm) (C–D are stereo pairs); E, scler-

ites from the medulla (scales 30 µm).

Description. Robust flattened branches, white cortex and pink to purple autozooidapertures. Numerous conical, semi-closed, autozooid polyp apertures uniformly/randomlydistributed throughout the branches (Fig. 39). Tiny siphonozooid apertures tightly closed,not observable to the naked eye. Medulla in the terminal branches with 6–7 major canals.Autozooid polyp tentacles with blunt ovals up to 0.1 mm long, regularly ornate with nota-ble conical rays (Fig. 40A–B) though different overall form in the paratype (Fig. 41A).


1014ZOOTAXA Surface sclerites mostly 7-radiates, and occasionally 8-radiates, with long (> 0.01 mm)

lobulated, smooth rays (Fig. 40D–G, 41B–D), averaging 0.065mm in length (±0.005 SD,n=10). Oval surface sclerites about 1.45 times longer than wider, averaging 0.045 mm inwidth (±0.005 SD, n=10). Medulla with highly ornate, forked, and irregular, spindles usu-ally less than 0.3 mm in length (Figs. 40H).

Morphological variation. Two characters exhibited a great deal of variation betweenthe holotype and paratype: the form of autozooid ovals and the medullar sclerites, the mostvariable character in Paragorgia spp. The medulla sclerites of the holotype of P.stephencairnsi sp. nov. have a great array of forked and bent forms. Since both examinedspecimens present the same kind of surface sclerites, which is the prevailing diagnosticcharacter in most Paragorgia spp., they can be considered as the same species until furtherspecimens are available.

Distribution. Pacific Ocean, off British Columbia, Canada, 350–490 m. Species comparisons. Externally, P. stephencairnsi sp. nov. is similar to P. yutlinux,

but the two can be differentiated using sclerite form. P. yutlinux has mostly 6-radiates andP. stephencairnsi 7-radiates.

Etymology: This species is named in honour of Dr. Stephen Cairns, one of the mostprolific coral taxonomists.

Genus Sibogagorgia Stiasny, 1937

Diagnostic characters. Paragorgiidae with scleritic medulla without large penetratingcanals, but with the main solenia around the subsurface/outer medulla (boundary canals)as a reticulate network (see details in Verseveldt, 1942). The network of canals can beobserved under the light microscope as a regularly reticulate and uniform mesh justbeneath the surface. Polyps without tentacular sclerites. Autozooid polyps uniform to ran-domly distributed along the branches.

Sibogagorgia weberi Stiasny, 1937(Fig. 42)

Sibogagorgia weberi Stiasny 1937: 80; Verseveldt 1942: 175.Not Suberia koellikeri: Nutting 1911: 13. Material examined. Holotype: ZMA Coel 03282 (Amsterdam), H-849, NIWA 3330,fragment, Siboga expedition st. 297, 10° 39’S–123° 40’E, 520 m, South of Timor, Indone-sia.

Diagnostic characters. Medulla sclerites smooth, barely ornate spindles exhibitingmost ornamentation at tips (Fig. 42F).


1014ZOOTAXA

FIGURE 42. Sibogagorgia weberi (Stiasny, 1937), holotype (ZMA Coel 03282): A–D, radiatesfrom the surface (scales 10 µm) (A–B are stereo pairs); E, intermediate sclerites from the subsur-face (scales 10 µm); F, sclerites from the medulla (scales 100 µm).

Description. Branching colonies probably larger than 200 mm in height, according toStiasny’s photographs (1937: plate IV). Terminal branches 3–7.5 mm in diameter (Stiasny1937). White to beige cortex. All sclerites colourless. Surface of the cortex with radiatesclerites (mostly 8-radiates) with star-like sub-radial ornamentation (Fig. 42A–E). Surfacesclerites up to 0.07 mm long averaging 0.064 (±0.007 SD, n=10) (Fig. 42A–D). Surfaceradiates about 1.64 times longer than wider, averaging 0.039 mm in width (±0.003 SD,n=10). Other intermediate forms between radiate and spindle present in the boundary layer(Fig. 42E). Medulla with smooth, barely ornate (most of ornamentation at tips) straightspindles usually less than 0.3 mm in length (Fig. 42F).

Morphological variation. The holotype specimen, the only known record of the spe-cies, presents slightly different types of sclerites at both the cortex and medulla. The sub-radiate ornaments vary from somewhat acute star-like tips (e.g., Fig. 42A–B) to smooth


1014ZOOTAXA and blunt (Fig. 42C–D). Medulla spindles are progressively less ornate towards the inner

medulla. Distribution. Only known for the type locality south of Timor, Indonesia, 520 m.

Sibogagorgia tautahi sp. nov.(Fig. 43)

Material examined.Holotype: NIWA 3327, H-847, J107, Z11229, 30° 1.891’S–178°48.061’W, “Giggenbach seamount”, 872–1086 m, 23 April 2002, (RV Tangaroa 0205/73,epibenthic sled).

Diagnostic characters. Sclerites from the medulla thick, blunt, short, mostly smooth,straight spindles that are ornate only at the tips, reaching usually less than 0.25 mm inlength (Fig. 43F).

Description. Holotype is a small (defectively preserved) colony (<40 mm long), 5 mmdiameter with 18 mm of base (Fig. 43G). Cortex pale brown. All sclerites colourless. Sur-face cortex with radiate sclerites (mostly 8-radiates) with star-like sub-radial ornamenta-tion (Fig. 43A–D). Surface sclerites up to 0.09 mm long averaging 0.08 (±0.007 SD,n=10). Surface radiates about 1.7 times longer than wide, averaging 0.047 mm in width(±0.004 SD, n=10). Other intermediate forms in the boundary layer include irregularlyenlarged radial ornamentation (Fig. 43E). Medulla with thick, blunt, short, mostly smooth,straight spindles, ornate only at tips, usually less than 0.25 mm in length (Fig. 43F).

Morphological variation. As in S. weberi, medullar spindles in S. tautahi sp. nov. areprogressively less ornate towards the inner medulla, achieving higher variation at theboundary layer (e.g., Fig. 43E). Some of the medullar sclerites are also quite robust andfatter than others (Fig. 43F).

Distribution. Only know from the Kermadec volcanoes seamount ridge, off the Bayof Plenty, New Zealand, 872–1086 m.

Species comparisons. S. tautahi sp. nov. has larger radiates than S. weberi, althoughoverall they are very similar, but those of S. tautahi are more profusely ornate (sub-radialornamentation). The diagnostic medullar spindles are shorter than those of S. weberi.

Etymology: The word tautahi means “solitary” in Maori and alludes to the sole smallspecimen of this species.


1014ZOOTAXA

FIGURE 43. Sibogagorgia tautahi sp. nov., holotype (NIWA 3327): A–D, radiates from the sur-face (scales 10 µm) (A–C are stereo pairs); E, intermediate sclerites from the subsurface/medulla(scales 10 µm); F, sclerites from the medulla (scales 100 µm); G. photograph of the entire colony(scale 20 mm).


1014ZOOTAXA Sibogagorgia dennisgordoni sp. nov.

(Figs. 44–46)

Material examined.Holotype: NIWA 3328, H-848, J237, Z8882, 37°01’S–176°43.1’E,“Jasons Hill”, 976–1129 m, August 1997 (col. J. Wills, FV Margret Phillipa 1024, bottomtrawl).

Paratype: NIWA 3329, P-1405, J246, Z9228, 36°41.7S–176°27.6E, western Bay ofPlenty slope, 820 m, 17 August 1998 (col. J. Wills, FV Margret Phillipa 1124/70, bottomtrawl).

Diagnostic characters. Cortex surface sclerites mostly 6- and 8-radiates with groovedand irregular sub-radial ornamentation (Figs. 45A–C, 46A–C). Medulla sclerites slim,long, bare, straight spindles, with club like tips, up to 0.4 mm in length (Figs. 45E, 46E).

FIGURE 44. Sibogagorgia dennisgordoni sp. nov., holotype (NIWA 3328) (calliper open at 20mm).

Description. Branched colonies up to 41 cm in height (holotype), terminal branches5–8 mm in diameter (Fig. 44). Cortex white to pale yellow. All sclerites colourless. Sur-face cortex with radiate sclerites (mostly 6- and 8-radiates) with grooved and irregular


1014ZOOTAXAsub-radial ornamentation (Figs. 45A–C, 46A–C). Surface sclerites up to 0.077 mm long

averaging between 0.06–0.07 (±0.004 SD, n=10, NIWA 3328; ±0.005 SD, n=10, NIWA3329). Surface radiates between 1.4–1.5 times longer than wider, averaging 0.04 mm inwidth (±0.003 SD, n=10, NIWA 3328; ±0.002 SD, n=10, NIWA 3329). Other intermediateforms between radiate and spindle present in the boundary layer (Figs. 45–46D). Medullawith slim, long, bare, straight spindles, with club like tips, up to 0.4 mm in length (Fig. 45–

46F).

FIGURE 45. Sibogagorgia dennisgordoni sp. nov., holotype (NIWA 3328): A–C, radiates from thesurface (scales 10 µm) (A–B are stereo pairs); D, intermediate sclerites from the subsurface/medulla (scales 10 µm); E, sclerites from the medulla (scales 100 µm).

Morphological variation. The irregular nature of the ornamentation of the surfaceradiates makes this character very variable. Nonetheless, all the characters are consistentin the two available specimens.


1014ZOOTAXA Distribution. Only known from the volcanic Kermadec Ridge, NE of the Bay of

Plenty, New Zealand, 820–1129 m.

FIGURE 46. Sibogagorgia dennisgordoni sp. nov., paratype (NIWA 3329): A-C, Radiates from thesurface (scales 10 µm) (A-B are stereo pairs); D, Intermediate sclerites from the subsurface/medulla(scales 10 µm); E, Sclerites from the medulla (scales 100 µm).


1014ZOOTAXASpecies comparisons. The diagnostic surface radiates of S. dennisgordoni sp. nov. are

so derived compared to those in other Sibogagorgia species that they more resemble thoseof some Paragorgia species. The other Sibogagorgia species do not have 6-radiate forms.Medullar sclerites, the other diagnostic character, are also quite modified, being more slen-der than in the other species. According to this character, S. dennisgordoni is closelyrelated to S. tautahi because both species have bare spindles with only ornaments at thesclerite tip but S. tautahi has shorter and fatter forms.

Etymology: The species is named in honour of Dr. Dennis Gordon, who has made out-standing contributions to marine diversity and who introduced me to the fascinating octoc-oral fauna of New Zealand.

Discussion

The deep-water bubblegum corals, Paragorgiidae, total 17 species, including the resultsfrom this paper. There were only five known species previously for Paragorgia (P.arborea, P. johnsoni, P. splendens, P. regalis [=dendroides], and P. coralloides) and justone of Sibogagorgia (S. weberi). As mentioned earlier, P. sibogae Bayer was not includedin this study due to the lack of a specimen: the original description was based entirely ondrawings from a sclerite slide preparation by Nutting without assigning a specimen as aholotype (Bayer 1993). In addition, since this study is based mostly on SEM images ofsclerites it would have been impossible to examine sclerites immersed in resin, as is thepresent condition of Nutting’s slide; besides, drawings of radiate sclerites are not sufficientto provide reliable species distinctions, as was noticed in this study by comparing SEMscans to previous drawings. The 11 new species included in this paper comprise 9 Para-gorgia species (P. alisonae, P. kaupeka, P. maunga, P. aotearoa, P. wahine, P. whero, P.yutlinux, P. stephencairnsi and P. tapachtli) and two Sibogagorgia species (S. tautahi andS. dennisgordoni). This study also highlighted two areas of endemism for bubblegum cor-als, corresponding to New Zealand and the Eastern Pacific (Canada and USA). NewZealand has six endemic species of Paragorgia (P. alisonae, P. kaupeka, P. maunga, P.aotearoa, P. whero, and P. wahine) and the two new species of Sibogagorgia (Fig. 2). P.yutlinux, P. stephencairnsi and P. tapachtli were collected in the Eastern Pacific (Fig. 1).The diversity of Paragorgiidae in New Zealand is remarkably high compared to otherPacific regions where even larger octocorals surveys have been undertaken such as Hawaii(e.g., Grigg & Bayer 1976) or Japan (e.g., Imahara 1996).

This study was primarily focused on the New Zealand fauna, of which there have beenrecent collections, particularly from habitats such as seamounts where bubblegum coralsseem to be very common. The diversity of Paragorgiidae species in New Zealand is thehighest for a single Economic Exclusive Zone. This trend is not entirely surprising giventhat other sessile colonial organisms such as bryozoans and stylasterids have a deep-waterdiversity “hotspot” in New Zealand (e.g., Gordon 1989; Cairns 1991). Unfortunately, most


1014ZOOTAXA of the new species from New Zealand are based on one or a few specimens, which suggest

that Paragorgiidae could be even more diverse in this region. Other groups of octocoralssuch as bamboo corals (Isididae) also have a diversity hotspot in Australasia particularlythe Mopseinae in Australia (Alderslade 1998) and the Keratoisidinae in New Zealand(Grant 1976; Smith et al. 2003; Sánchez in prep.). Consequently, the great diversity of Par-agorgiidae in New Zealand is not surprising but a trend found in other benthic sessileorganisms including octocorals (Sánchez & Rowden accepted). The new material from theEastern Pacific, deposited in the Smithsonian Institution, revealed also another likelyendemic region for Paragorgiidae. It would be valuable to have access to additional mate-rial from both regions, particularly seamounts, which could provide additional new spe-cies.

The phylogenetic relationships amongst Paragorgiidae obtained with morphologicalcharacters reflected the evolution of surface sclerites, which are the most informative char-acters. Surface sclerites in gorgonian corals, e.g., octocorals with a scleroproteinaceousaxis, modify colony flexion, thereby helping the compressibility of the outer cortex whenthe sclerites’ enlarged asymmetrical sides contact each other (Lewis & Von Wallis 1991).Analogously, one clade of Paragorgia species exhibits asymmetrical surface sclerites (P.maunga, P. kaupeka, P. coralloides, P. tapachtli and P. regalis). The function of sclerites inparagorgiids is unknown, but since the species have no axial skeleton other than the scler-ites themselves, they should definitely be under high biomechanical stress and hence underselection by hydrodynamic regimes. Consequently, it is possible that apparently homolo-gous forms in those sclerites could evolve in parallel, creating homoplasy, as has beenobserved in the Plexauridae (Sánchez unpublished). Even so, scleritic characters providesufficient support to discern and distinguish all the Paragorgiidae species, construct a spe-cies key, and provide phylogenetic hypotheses. The presence of homoplasy in surfacesclerites needs further scrutiny using independent characters.

Bubblegum corals have one of the most intriguing geographical patterns for deep-water corals. P. arborea in particularly has an apparently cosmopolitan but highly discon-tinuous distribution (e.g., Fig. 1). P. arborea seems widely distributed at subpolar oceansat depths below 200 m but no records have been found at equatorial latitudes (Broch 1957;Grasshoff 1979). This study examined a specimen from Alaska and no major morphologi-cal differences with New Zealand specimens were found. In fact, the same diagnostic char-acters for the species were consistently found at either locality, though the Alaskaspecimen had overall smaller sclerites. New Zealand specimens of P. arborea include thedeepest record for the species, 1525 m (NIWA 3311), whereas it has been found above1330 m at northern latitudes (Tendal 1992). Unfortunately, and intriguingly, the dispersalmechanisms of paragorgiids between latitudes are unknown, as is whether they are geneti-cally different species, or if there are connecting populations at equatorial latitudes (whichmay require sampling at greater depths). Since P. arborea appears basal to the rest of thespecies, as well as the most widely distributed, it would be also interesting to test if it is a


1014ZOOTAXAphylogeographical pattern for the group, wherein the most basal species retain the ances-

tral distribution. Nonetheless, P. arborea seems to be exceptional within the Paragorgiidaebecause most species have a more restricted distribution. There also seems to be a patternof specialization of surface sclerites for derived species of Paragorgia (e.g., P. yutlinuxwith only 6-radiates) whereas P. arborea, presumably basal, contains 6- to 8-radiates,which could be the ancestral state. There seems to be a single trans-Pacific species, viz P.regalis, and likewise in the Atlantic with P. johnsoni (e.g., Fig. 1) but it is clear that noother species is as widespread as P. arborea. There are cases of morphological sister spe-cies such as P. johnsoni and P. aotearoa that correspond to the Atlantic and Pacific respec-tively but the phylogenetic relationships of the other species suggest that mostParagorgiidae diversity and speciation took place in the Pacific Ocean. The nine speciesfrom New Zealand were mostly collected between 800 and 1000 meters, which suggests asimilar habitat requirement for most bubblegum corals found there. Although most of thespecies were found north of New Zealand, some species were found both north and south(e.g., P. alisonae and P. whero). Perhaps more sampling to the south is necessary to ascer-tain if the endemic species are found throughout New Zealand. It is important to mentionalso that two of the new species were collected off the Chatham Islands exclusively (P.aotearoa and P. wahine), where intensive fishing takes place. A limited range in tempera-ture, allowing a large depth range according to latitude, seems to be the key parameter forthe distribution for Paragorgia arborea (Tendal 1992). However, there are some speciesnear the equator, such as P. coralloides and P. johnsoni, and it is premature to give conclu-sions on habitat and distributional preferences for the Paragorgiidae prior to more system-atic sampling.

Acknowledgments

This study was funded totally by a postdoctoral research fellowship at the National Insti-tute of Water and Atmospheric Research, New Zealand. I greatly acknowledge DennisGordon who encouraged me to study of bubblegum corals and the fascinating NewZealand octocoral fauna. Stephen Cairns (Smithsonian Institution) provided valuable helpand photographs of the specimens of the Eastern Pacific. Collaboration, advice and helpfrom Malcolm Clark, Ashley Rowden, Megan Oliver, Di Tracey, Dean Stutter, Anne-NinaLoerz (NIWA Invertebrate Collection), Alison R. Acosta de Sánchez, Jim Boutillier, R.Van Soest. Kay Card (Industrial Research), Allen Andrews and P. Smith are greatlyacknowledged. Comments from Phil Alderslade (Darwin, Australia) and an anonymousreviewer greatly improved the contents of this paper. The University of the Andes (Depar-tamento de Ciencias Biológicas, Facultad de Ciencias, Bogotá, Colombia) providedresources and facilities to finish this study. The British Museum (Natural History),Amsterdam Museum, and Smithsonian Institution kindly provided valuable and historicaloctocoral type material.


1014ZOOTAXA References

Alderslade, P. (1998) Revisionary systematics in the gorgonian family Isididae, with descriptions ofnumerous new taxa (Coelenterata: Octocorallia). Records of the Western Australian Museum,55, 1–359.

Allemand, D. (1993) The biology and skeletogenesis of the Mediterranean Red Coral: A review.Precious Corals & Octocoral Research, 2,19–39.

Bayer, F. M. (1956a) Octocorallia, In: Moore, R.C. (Ed.), Treatise on Invertebrate PalaeontologyPart F. Coelenterata. Lawrence, Kansas: Geological Society of America and University ofKansas Press, 163–231

Bayer, F. M. (1956b) Descriptions and redescriptions of the Hawaiian octocorals collected by theU.S. Fish Commission steamer "Albatross" (2. Gorgonacea: Scleraxonia). Pacific Science, 10,1, 67–95.

Bayer, F.M. (1961) The shallow water Octocorallia of the West Indian region. Studies on the Faunaof Curaçao, 12, 1–373.

Bayer, F.M. (1964) A new species of the Octocorallian genus Paragorgia trawled in Florida watersby R.V. Gerda. Zoologische Mededelingen, Leiden, 39, 526–532.

Bayer, F.M. (1992) The Helioporacean Octocoral Ephiphaxum, recent and fossil: a monographiciconography. Studies in Tropical Oceanography, 15, 1– 76.

Bayer, F.M. (1993) Two new species of the gorgonacean genus Paragorgia (Coelenterata: Octocor-allia). Precious Corals & Octocoral Research, 2, 1–40.

Broch, H. (1912) Die Alcyonarien des Trondhjemsfjordes. II. Gorgonacea. Kongelige NorskeVidenskabers Selskabs Skrifter, 2, 1–48.

Broch, H. (1957) The Northern octocoral, Paragorgia arborea (L.), in sub-antarctic waters. Nature,179, 1356.

Cairns, S.D. (1991) The marine fauna of New Zealand: Stylasteridae (Cnidaria: Hydroida). NewZealand Oceanographic Institute Memoir, 98, 1–179.

Buhl-Mortensen, L. & Mortensen, P. B. (2004) Crustaceans associated with the deep-water gorgon-ian coral Paragorgia arborea (L., 1758) and Primnoa resedaeformis (Gunn., 1763). Journal ofNatural History, 38, 1233–1247.

Fu, J., & Murphy, R.W. (1999) Discriminating and locating covariance: an application of permuta-tion tail probability (PTP) analyses, Systematic Biology, 48, 380–395.

Gordon, D.P. (1989) The marine fauna of New Zealand: Bryozoa: Gymnolaemata (Cheilostomida:Ascophorina) from the western South Island continental shelf and slope. New Zealand Ocean-ographic Institute Memoir, 97, 1–158.

Grant, R. (1976) The marine fauna of New Zealand: Isididae (Octocorallia: Gorgonacea) from NewZealand and the Antarctic. New Zealand Oceanographic Institute Memoir, 66, 1–56.

Grasshoff, M. (1979) Zur bipolaren verbreitung der Oktokoralle Paragorgia arborea (Cnidaria:Anthozoa: Scleraxonia). Senckenbergiana maritime, 11, 3/6, 115–137.

Grasshoff, M. (1979) Neubeschreibung der Oktokoralle Paragorgia johnsoni Gray, 1862 (Cnidaria:Anthozoa: Scleraxonia). Senckenbergiana biologica, 60, 427–435.

Gray, J.E. (1862) Notice of a second species of Paragorgia discovered in Madeira by Mr. JamesYate Johnson. Annals and Magazine of Natural History, 10, 125–126.

Grigg, R.W. & Bayer, F.M. (1976) Present knowledge of the systematics and zoogeography of theorder Gorgonacea in Hawaii. Pacific Science, 30, 167–175.

Heifetz, J. (2002) Coral in Alaska: distribution, abundance, and species associations. Hydrobiolo-gia, 471, 19–28.

Imahara, Y. (1996) Previously recorded octocorals from Japan and adjacent seas. Precious Corals& Octocoral Research, 4–5, 17–44.

Kitching, I. J., Forey, P. L., Humphries, C. J. & Williams, D. (1998) Cladistics: theory and practice


1014ZOOTAXAof parsimony analysis. 2nd Edition. Systematics Association Special Volumes, Oxford Univer-

sity Press, pp 248.Koren, J. & Danielssen, D. C. (1877) Nye Alcyonider, Gorgonidr og Pennatulider tilhørende

Norges Fauna. Bergens Museum, Bergen, John Griegs Bogtrykkeri, 38 pp.Koslow, J.A., Gowlett-Holmes, K., Lowry J.K., O'Hara T., Poore, G.C.B. & Williams, A. (2001)

Seamount benthic macrofauna off southern Tasmania: Community structure and impacts oftrawling. Marine Ecology Progress Series, 213, 111–125

Krieger, H.J. (1993) Distribution and abundance of rockfish determined from a submersible andbottom trawling. Fisheries Bulletin, 91, 87–96.

Krieger, K. J. & Wing, B. (2002) Megafauna associations with deepwater corals (Primnoa spp.) inthe Gulf of Alaska. Hydrobiologia, 471, 83–90.

Lewis, J. C. & Wallis, von E. (1991) The function of surface sclerites in gorgonians (Coelenterata,Octocorallia). Biological Bulletin, 181, 275–288.

Linnaeus, C. (1758) Systema Naturae, Editio Decima, reformata 1, Holmiae.Nixon, K. C. (1999) Winclada (BETA) ver. 0.9.9. Published by the Author, Ithaca, NY, 7 pp.Nutting, C.C. (1911) The Gorgonacea of the Siboga Expedition VIII. The Scleraxonia. Siboga-

Expedition Monographs, 13b, 1–62.Nutting, C.C. (1912) Descriptions of the Alcyonaria collected by the U.S. Fisheries steamer "Alba-

tross," mainly in Japanese waters, during 1906. Proceedings of the United States NationalMuseum, 43, 1–104.

Sánchez, J.A. (2001) Systematics of the Southwestern Caribbean Muriceopsis Aurivillius (Cni-daria: Octocorallia) with description of a new species. Bulletin of the Biological Society ofWashington, 10, 160–180.

Sánchez, J.A., & Cairns, S.D. (2004) An unusual new gorgonian coral from the Aleutian Islands,North Pacific, Alaska. Zoologische Mededelingen, Leiden, 78, 265–274.

Sánchez, J.A., & Rowden A. A. (accepted) Octocoral diversity on New Zealand seamounts. Pro-ceedings of the XI International Coral Reef Symposium. Okinawa, Japan.

Sánchez, J.A., Lasker, H.R. & Taylor, D.J. (2003) Phylogenetic analyses among octocorals (Cni-daria) according to mitochondrial and nuclear DNA sequences (lsu-rRNA 16S, and ssu-rRNA18S) support two convergent clades of branching gorgonians. Molecular Phylogenetics & Evo-lution, 29, 1, 31–42. (DOI:10.1016/S1055-7903(03)00090-3)

Sánchez, J.A., Lasker, H.R., Nepomuceno, E. G., Sánchez, J. D. & Woldenberg, M. J. (2004)Branching and Self-Organization in Marine Modular Colonial Organisms: a Model. The Amer-ican Naturalist, 163, 3, E24–39. (DOI: 0003-0147/2004/16303-20458).

Smith, P. J., S. M. McVeagh, J. T., Mingoia, & S.C. France (2003) Mitochondrial DNA sequencevariation in deep-sea bamboo coral (Keratoisidinae) species in the southwest and northwestPacific Ocean. Marine Biology (in press).

Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and other methods). Ver-sion 4.0b10. Sunderland: Sinauer Associates.

Stiasny, G. (1937) Die Gorgonacea der Siboga-Expedition. Supplement II, Revision der Scleraxoniamit ausschluss der Melitodidae und Coralliidae. Siboga-Expedition Monographs, 13b, 1–138.

Tendal, O.S. (1992) The North Atlantic distribution of the octocoral Paragorgia arborea (L., 1758)(Cnidaria, Anthozoa). Sarsia, 77, 213–217.

Thomson, J.A. & Henderson W.D. (1906) An account of the alcyonarians collected by the RoyalIndian Marine Survey Ship Investigator in the Indian Ocean. I. The Alcyonarians of the deepsea. The Trustees of the Indian Ocean Museum, Calcutta, 132 pp.

Verrill, A. E. (1922) Part G: Alcyonaria and Actiniaria. Report of the Canadian Arctic Expedition1913–18, 8, 1–164.

Verseveldt, J. (1940) Studies on Octocorallia of the families Briareidae, Paragorgiidae andAnthothelidae. Temminckia, 4, 1–142.


1014ZOOTAXA Verseveldt, J. (1942) Further studies on Octocorallia. Zoologische Mededelingen, Leiden, 24, 159–

186.Watling, L. & Norse, E.O. (1998) Disturbance of the seabed by mobile fishing gear: a comparison

to forest clear cutting. Conservation Biology, 12, 1180–1197.

Zootaxa, Cnidaria, Octocorallia, Paragorgiidaedocs.niwa.co.nz/library/public/1877407194.pdf · Systematics of the bubblegum corals (Cnidaria: Octocorallia: Paragorgiidae) with description

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