European Journal of Taxonomy 68: 1-26 ISSN 2118-9773

http://dx.doi.org /10.5 852/ejt.2013.68 www.europeanjoumaloftaxonomy. eu

2013 • Sabine Stohr, Emilie Boissin & Thierry B. Hoareau

This work is licensed under a Creative Commons Attribution 3.0 License.

Research article

um:lsid:zoobank.org:pub:AD45942C-4285-4BAD-9728-EB41EE22F226

Taxonomic revision and phylogeny of the Ophiocoma brevipes group

(Echinodermata, Ophiuroidea), with description of a new subgenus

(Breviturma) and a new species

Sabine STOHR', Emilie BOISSIN" & Thierry B. HOAREAU"

^Swedish Museum of Natural History, Department of Zoology, Box 50007, 104 05 Stoekholm, Sweden

Email: [email protected] (eorresponding author)

^’^Eaboratoire ECOMAR, LJniversite de Ea Reunion, BP 7151, 97715 Saint-Denis, Ea Reunion, Franee

2USR3278-CRIOBE-CNRS-EPHE, Eaboratoire d’Exeellenee « CORAIE »,

LJniversite de Perpignan-CBETM, 58 rue Paul Alduy, 66860 Perpignan Cedex

^Email: [email protected]

^Moleeular Eeology and Evolution Programme, Department of Geneties, University of Pretoria,

Private bag X20, Hatfield, Pretoria 0028, South Afriea

^Email: [email protected]

^ um:lsid:zoobank.org:author:412800EB-AACE-4313-9810-61F89B740405

^ um:lsid:zoobank.org:author:B35FF544-AE44-43B2-BlCA-3769A281F386

^ um:lsid:zoobank.org:author:44578720-231E-4D99-8F38-2A94B7C4E4F2

Abstract. The taxonomy of the genus Ophiocoma was last revised by Devaney in 1970. Reeent diseoveries

of new speeies and re-instatement of previously synonymized names suggest that we still do not fully

understand the speeies limits in this genus. A reeent biodiversity survey of the SW Indian Oeean shallow

reefs strongly suggested an unreeognised speeies in the genus, elosely related to O. brevipes!O. dentata.

This study examined both the moleeular phylogenetie relationships and the morphologieal eharaeteristies

of several speeies in the genus in order to eharaeterise the unreeognised speeies. The foeal speeies elusters

with O. brevipes, O. dentata, O. doederleini within a monophyletie elade supported by moleeular data

for the first time. The name Breviturma subgen. nov. is proposed for this elade, previously known as

brevipes group. Type material of nominal speeies that have been synonymized with O. dentata was

examined and re-assessed. Ophiocoma marmorata proved not eonspeeifie with O. dentata. A rarely

used eharaeter, dorsal dise granule density, was tested and showed differenees between the examined

speeies at similar sizes. In eombination with eolour pattern, dise granule density, arm spine sequenee

and maximum dise size, the new speeies was delimited morphologieally and deseribed as Ophiocoma

krohi sp. nov.

Key words, morphology, Ophiocoma krohi, Breviturma, mitoehondrial DNA, eryptie biodiversity.

Stohr S., Boissin E. & Hoareau T.B. 2013. Taxonomic revision and phylogeny of the Ophiocoma brevipes group

(Echinodermata, Ophiuroidea), with description of a new subgenus {Breviturma) and a new species. European

Journal of Taxonomy 68: 1-26. http://dx.doi.org/10.5852/ejt.2013.68

European Journal of Taxonomy 68 : 1-26 ( 2013 )

Introduction

The history of the taxonomy of the genus Ophiocoma L. Agassiz, 1835 is eonvoluted and eonfused.

Devaney (1970) resolved some of the eonfusion using external and internal eharaeters sueh as spine

sequenees along the arm, eolour pattern, and the shape of the dental plate, to delimit four groups,

namely the brevipes, scolopendrina and pumila groups earlier proposed by Clark (1921), and the

pica group (Devaney 1970). After 1970, few new speeies of Ophiocoma have been deseribed, namely

O. paucigranulata Devaney, 1974 from the Caribbean and O. aegyptiaca Soliman, 1991 from Egypt.

Also, O. endeani Rowe & Pawson, 1977 was proposed as a replaeement name for O. alternans Endean,

1964 (a homonym) by Rowe & Pawson (1977). Ophiocoma similanensis Bussarawit & Rowe, 1985

was synonymized with O. erinaceus Muller & Trosehel, 1842 by Priee & Rowe (1996). Reeently a

new speeies O. cynthiae Benavides-Serrato & O’Hara, 2008 was deseribed and O. schoenleini Muller

& Trosehel, 1842 was reinstated as a valid speeies (Benavides-Serrato & O’Hara 2008). Both are elose

to O. erinaceus and have previously been mistaken for it, as was suggested by an earlier moleeular

study (O’Hara et al. 2004). Although Ophiocoma is a well studied genus, eonsisting of eomparatively

large, eonspieuous, abundant and easily aeeessible tropieal shallow water speeies, it elearly still holds

unreeognized diversity. Currently, 22 speeies are aeeepted in the genus (Stohr & O’Hara 2013).

In a reeent large-seale marine biodiversity survey in the Southwestern Indian Oeean (BIOTAS), we

identified several speeimens as belonging to the brevipes group based on morphology. Aeeording to

Devaney (1970), the brevipes group eurrently ineludes the three speeies Ophiocoma brevipes Peters,

1851, O. dentata Muller & Trosehel, 1842 and O. doederleini de Eoriol, 1899. In our material, we

found speeimens belonging to these three speeies and a fourth unknown speeies was suspeeted and

eonfirmed by subsequent moleeular analysis. They formed a distinet genetie lineage when eompared to

other available speeies of Ophiocoma (Hoareau et al. 2013). Currently (Devaney 1970; Stohr & O’Hara

2013), the following nominal speeies are regarded as synonyms of the speeies in the brevipes group:

O. brevispinosa Smith, 1876 = O. brevipes', O. insularia Eyman, 1862, O. marmorata Marktanner-

Tumeretseher, 1887, O. ternispina v. Martens, 1870 and O. variegata Smith, 1876 = O. dentata. The

name Ophiopeza danbyi Farquhar, 1897 was ineluded in the synonymy of O. brevipes by Clark (1915),

but not by Devaney (1970). The newly reeognized lineage may either belong to a new, not yet deseribed

speeies, or it may eorrespond to one of the synonymized speeies.

Speeimens of the brevipes group are eharaeterized by similar numbers of arm spines on eaeh side of the

arm segments beyond the dise, four to six arm spines on eaeh side of proximal arm segments, a dental

plate that is roughly twiee as long as wide, with wide septum dividing the tooth foramina, broadly

oval dorsal arm plates, densely paeked, small and spherieal dise granules that eover the entire ventral

interradius, and two tentaele seales along most of the arm (Devaney 1970). Aeeording to Devaney (1970),

the eomparatively short dental plate with small region for the tooth papillae is an important eharaeter to

distinguish the brevipes group from the other subgroups of Ophiocoma. Moreover, Benavides-Serrato &

O’Hara (2008) found that subtle differenees in eharaeters sueh as the extent of granulation on the ventral

dise, tube foot eolour and shape of the dental plate had previously been overlooked or misinterpreted

as intraspeeifie variation in speeies of the scolopendrina group. These eriteria might be informative to

eharaeterise the speeimens of the newly found lineage in the brevipes group.

This study aims to deseribe that unidentified speeies in the brevipes group, using both moleeular methods

and morphologieal eharaeters. Using two mitoehondrial loei, we eonstrueted a phytogeny to assess the

moleeular status of the brevipes group relative to the scolopendrina group and to evaluate the speeies

relationships within the brevipes group. In addition, we identified a new eharaeter, dorsal dise granule

density, to distinguish between the speeies in the brevipes group and re-evaluate type material of the

studied speeies.

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

Material and methods

Brittle stars of the genus Ophiocoma were eolleeted at La Reunion Island and off Madagasear (Nosy-

Be Island). Speeimens of O. doederleini from La Reunion and Polynesia were provided by G. Paul ay

from the eolleetions of the Florida Museum of Natural History. Total genomie DNA was extraeted

from a pieee of arm following the DNeasy protoeol (Qiagen). COI sequenees were already available

for most of the speeies (Hoareau et al. 2013; Supplementary file: KC759738-KC759923) exeept for

O. dentata and O. doederleini. A portion of the COI bareoding gene and of the I6S ribosomal gene

were amplified using eehinoderm-speeifie hybrid primers (Hoareau & Boissin 2010) and universal

primers (Palumbi 1996), respeetively. We followed the PCR eonditions deseribed in these studies.

The amplieons were then sequeneed using BigDyeTerminator (Applied Biosystems, Foster City, CA,

USA) eyele sequeneing reaetions, and eleetrophoresed on an ABI 3730x1 DNA Analyzer (Applied

Biosystems) at the Interdiseiplinary Center for Bioteehnology Researeh (University of Florida). We

aligned the sequenees using mafft 6 online I http://mafft. ebre.ip/alignment/server/ I with all the default

settings. All the sequenees are available in GenBank (see Supplementary file). From the eoneatenated

datasets, a Neighbour Joining phylogenetie tree was reeonstrueted using Mega5.05 (Tamura et al. 2011)

applying 1000 bootstraps and the Kimura-2-Parameters model of substitution. Additionally, we used

maximum likelihood inferenee using Mega. Beeause we used two mitoehondrial loei, we estimated

a single substitution model for the eoneatenated dataset using Mega (GTR + gamma + invariant site

model). Support for eaeh node was assessed with 200 bootstrap replieates. To overeome potential

eomplieations introdueed by alignment gaps, we applied the Partial-Deletion option available in Mega

in both reeonstruetion methods. Sequenees of Ophiocoma erinaceus and O. scolopendrina (Lamarek,

1816) from the scolopendrina group were used as outgroup.

After DNA extraetion, some of the individuals were used for morphologieal examination. Sinee the

new speeies is smaller than the others, we seleeted eomparative material from other speeies at similar

size (11-13 mm dise diameter), depending on availability. Two of the largest speeimens of Ophiocoma

sp. nov. at 11 mm and 13 mm dise diameter (dd) and a speeimen eaeh of O. doederleini (13 mm dd),

O. brevipes (13 mm dd) and O. dentata (11 mm dd) were seleeted for seanning eleetron mieroseopy

(SEM). These speeimens were first photographed with a digital eamera. Then they were treated with

diluted (1:1) household bleaeh (NaOCl) to remove the epidermis. They were mounted on aluminium

stubs with spray glue, dorsal side up, and examined in a Hitaehi FE-S4300 seanning eleetron mieroseope.

Then the glue was dissolved with butyl aeetate and the animals were remounted, ventral side up, and

examined by SEM again. Finally, the speeimens of O. doederleini (UF5318), O. brevipes (SMNH-

133230), O. dentata (URUN2009-11047) and the 13 mm dd Ophiocoma sp. nov. (SMNH-Type 8535)

were disartieulated in eoneentrated bleaeh. The skeletal elements were washed in tap water, mounted on

stubs and examined by SEM.

Whole animals were measured using a digital ealliper, smaller parts were measured on the SEM images

using the software ImageJ v.L47e (Rasband 1997-2012). Dise diameter is defined as the distanee from

the distal edge of a pair of radial shields (or if eoneealed the dise edge atop the arm) to the edge of the

opposite interradius. Densities of dise granules were measured by overlaying a square frame of 0.5 mm

side length (0.25 mm^) to the image in ImageJ and eounting all eomplete granules inside the square. This

was repeated at least three times in different plaees of the image, but always in the eentral part of the

dise to avoid the often bulging, more sparsely granulated dise periphery. The numbers were multiplied

by four to give densities for 1 mm^ for easier eomparability with previously published values. Vertebrae

were measured in ImageJ aeross their greatest width and height; oral plates (jaws) were measured by

fitting an ellipse around their irregular shape and ealeulating the major (here height) and minor (here

width) axes. Arm spines were eounted on 2-5 arms of eaeh speeimen and when the numbers at a segment

varied between arms the less eommon number is given in parentheses. The taxonomy follows Stohr &

European Journal of Taxonomy 68 : 1-26 ( 2013 )

O’Hara (2013). The morphological terminology follows Stohr et al. (2012). Field numbers, museum

registration numbers and GenBank accession numbers are listed in the Supplementary file.

Abbreviations

Acronyms of institutions and repositories

NHMW = Natural History Museum Wien (Vienna)

SMNH = Swedish Museum of Natural History

UF, FLMNH = Florida Museum of Natural History

URUN = Universite de La Reunion

ZMB = Zoological Museum Berlin

Other abbreviations

DAP =

dorsal arm plate

disc diameter

maximum likelihood

neighbour j oining

spm(s) =

specimen(s)

W:L =

width: length

Breviturma

subgen. nov.

scolopendrina

group

Fig. 1. Phylogenetic reconstruction of Ophiocoma {Breviturma) subgen. nov. (two species of the

scolopendrina group as outgroup), estimated by maximum likelihood and neighbour-joining analysis

of concatenated sequences of the mitochondrial cytochrome c oxidase subunit 1 and rDNA 16S (in total

1125 bp). Values at branches indicate bootstrap values for maximum likelihood (above branches) and

neighbour-joining (below branches). Specimen codes are resolved in the Supplementary file.

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

Results

The eoneatenated sequenees of the 17 speeimens eontained 1125 sites (1069-1119 bp) of whieh 303

(26.9 %) were parsimony informative. For most of the speeimens the eomplete target sequenee was

obtained. For the speeimens of O. doederleini, 614 bp were reeovered from the original 659 bp. The

16S sequenees varied depending on the presenee of multiple indels (450-456 bp). For the COl loeus, no

indels, distinet double pies or stop eodons were deteeted whieh rule out the presenee of pseudogenes.

Only three amino-aeids showed variation in the COl sequenees; one site involved a ehange from Serine

to Alanine in all the speeimens of O. krohi sp. nov.

Most of the nodes in the phylogeny reeeived a good bootstrap support (ML>95 and NJ>90). The ML

and NJ phylogenetie trees were eongruent and eonfirmed, for the first time, a moleeular basis for the

elustering of speeies from the brevipes group (100/100 for ML and NJ respeetively), relative to the

two speeies belonging to the scolopendrina group. The trees also revealed four highly supported taxa

(100/100) within the brevipes group, matehing the speeies O. brevipes, O. dentata, O. doederleini and

a new speeies (Fig. 1). Ophiocoma doederleini had a deeper branehing in the tree relative to the three

other speeies. The elade eonsisting of Ophiocoma sp. nov., O. brevipes and O. dentata was supported

(bootstrap values 97/91.9). However, the relationships within the elade were unelear sinee the elade

eonsisting of O. brevipes and O. dentata was weakly supported (56/65.5).

Comparison with the type material eonfirmed our identifieations of O. brevipes, O. dentata and

O. doederleini. The morphologieal examinations also supported the existenee of a fourth speeies in the

brevipes group. The most important eharaeters were dorsal dise granule density and arm spine numbers,

summarized in Table 1. Colour pattern was also an important speeies delimiting eriterion (Fig. 2).

Finally, the speeimens of Ophiocoma krohi sp. nov. were on average signifieantly smaller than the three

other speeies of the brevipes group (Kruskal-Wallis test, K=9.25\A, d.f=3, P =0.0261). This result

eonfirms previous observations based on examinations of speeimens in the eolleetions of FLMNH from

a large geographie range in the Paeifie and Indian Oeeans (Okinawa, Tuamotu Islands, Mariana Islands,

Masearene Islands, Samoa Islands, Soeiety Islands, Niue Island, Hawaiian Islands).

The brevipes group is well supported as a monophyletie group and below proposed as a subgenus of

Ophiocoma. The fourth speeies is deseribed below as new.

Class Ophiuroidea Gray, 1840

Order Ophiurida Muller & Trosehel, 1840

Family Ophioeomidae Ljungman, 1867

Genus Ophiocoma L. Agassiz, 1835

Diagnosis

Typieally large, eonspieuous, shallow water tropieal brittle stars. Dorsal dise eompletely eovered with

granules, extending onto the ventral side to varying degree. Strong, ereet, smooth and generally solid

(exeept O. pusilla Broek, 1888) arm spines. A eluster of tooth papillae on the lower part of the dental

plate; large reetangular teeth. Adoral shields eonfined to lateral edges of oral shield, not meeting. Abradial

musele surfaee of oral plates well developed with a series of horizontal grooves and ridges. Lateral arm

plates short, eurved, with ventral noteh for tentaele foot, spine artieulations sigmoidal.

European Journal of Taxonomy 68 : 1-26 ( 2013 )

Table 1. Morphometric data for the species of Ophiocoma (Breviturma) subgen. nov. Type material from

Zoological Museum Berlin (ZMB), Swedish Museum of Natural History (SMNH) and Natural History

Museum Vienna (NHMW). Specimen NHMW 104.66a is regarded as the holotype of O. marmorata, the

other specimens of that lot most likely belong to a different species. DD, disc diameter, MNHN, Museum

national d’Histoire naturelle (Paris), UF, Florida Museum of Natural History, URUN, University of La

Reunion.

Species

Specimen

DD [mm]

Granules

mm'^

5 spines at

segments

4 spines beyond

segment 15

Ophiocoma brevipes

ZMBEch 961

12-13

5-9

yes

ZMB Ech 962

3-14

yes

ZMB Ech 963

5-11

yes

ZMB Ech 4660

17-18

6-12

yes

ZMB Ech 4660

6-11

yes

ZMB Ech 4660

6-17

yes

SMNH-133230

240-280

6-10

yes

URUN 2013-11188

19.7

176-188

6-18

yes

URUN 2008-04774

13.1

148-168

6-10

yes

URUN 2008-05819

13.5

180-200

6-11

yes

Ophiocoma dentata

ZMBEch 931

18-19

none

URUN 2009-11047

216-244

none

SMNH-133232

23.7

44-64

(8, 9)

URUN 2007-00380

52-68

none

Ophiocoma ternispina

ZMB Ech 1815

none

Ophiocoma insularia

SMNH-Type-5244

17.6

96-120

7-9

yes

Ophiocoma marmorata

NHMW 10.466a

8.7

none

yes

NHMW 10.466b

6.3

none

yes

NHMW 10.466c

6.1

none

yes

NHMW 10.466d

5.2

6-7

yes

Ophiocoma krohi sp. nov.

SMNH-Type-8531

6.8

152

6-7

MNHN-IE-4303

125

none

SMNH-Type-8532

9.6

116-120

none

MNHN-Type-8533

140-164

6-9

yes

UF-13938

10.3

112-120

7-9

SMNH-Type-8534

132-136

6-7

yes

URUN 2009-11108

11.3

112-120

none

MNHN-IE-4301

11.8

72-96

5-8

yes

MNHN-IE-4300

12.7

104

none

SMNH-Type-8535

104

6-7

yes

MNHN-IE-4302

96-104

none

SMNH-Type-8536

15.5

64-96

none

yes

Ophiocoma doederleini

UF5318

200-224

6-7

yes

UF6512

88-96

6-9

yes

UF 6634

60-64

6-10

yes

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

Subgenus Breviturma subgen. nov.

um:lsid:zoobank.org:aet:DA2BAD13-QEQ3-45B3-9D7A-78796329BC3C

Diagnosis

Subgenus of Ophiocoma, previously known as brevipes group. Charaeterized by equal (non-alternating)

numbers of arm spines; dise granules uniform in size, low and blunt; dorsal arm plates rounded, almost

oval (in adult speeimens, more rounded triangular in younger individuals); dental plate about twiee as

long as wide and thus the shortest in the genus, with tooth papillae region oeeupying 25 % of the dental

plate length.

Etymology

The name Breviturma refers to the original name brevipes group, eombined with turma = troop, here

meaning group.

Type species

Ophiocoma brevipes VqXqvs, 1851.

Included species

Ophiocoma Boederleini, O. brevipes, O. dentata and O. krohi sp. nov.

Ophiocoma {Breviturma) doederleini de Loriol, 1899

Figs 2A-C, 3

Ophiocoma doederleini de Loriol, 1899: 30, pi. 3 fig. 2.

Diagnosis

Speeies of Ophiocoma {Breviturma) subgen. nov. eharaeterized by strongly dark and light banded

(annulated) arm spines; slender arm spines, the longest up to three segments long; up to five arm spines

on few segments from segment 6; variable in dorsal dise eolour pattern (retieulating or spotted); density

of dorsal dise granules reaehes more than 200 mm ~^. Maximum size at least 26 mm aeeording to the

studied material.

Material examined

UF5318

Collecting data. Polynesia, Moorea, 17°28’55” S, 149°50’32” W, fore reef between Cook’s and

Opunohu Bays, outer reef slope, under roeks, partly in sand, 10-15 m, by hand, eolleetors C. MeKeon,

C. Meyer, G. Paulay, 19 Jul. 2006, 1 spm.

Morphological data. 13 mm dd. Granule density on light image 180-200 mm ~^. Arm spine sequenee

3, 3, 3, 3, 4, 5, 5, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 3, 3. Dorsal dise dark brown

with irregular white patehes, with a net-like retieulating darker brown pattern (Fig. 2A, B). Dorsal arms

dark brown with widely spaeed white bands with brown spots. Ventral dise white with brown spots.

Oral frame white, oral shields with brown spots. Ventral arms with long white and brown bands, brown

spots on the white bands. Arm spines strongly annulated with white and brown rings, long and slender,

dorsalmost spine barely two segments long, middle spines longer than two segments.

Dorsal dise granule diameter 47-51 pm, granule density 224 mm'^ (Fig. 3A, C). Third spine on proximal

segments 2.3 mm long; proximal DAPs 1.74-1.8 mm wide (Fig. 3B). Oral shields slightly wider than long;

5-6 tooth papillae, 5 lateral oral papillae at eaeh jaw edge, outer papilla widest (Fig. 3D). Dental plate

European Journal of Taxonomy 68 : 1-26 ( 2013 )

Fig. 2. Colour variations in species of Ophiocoma (Breviturmd) subgen. nov. A-C. Ophiocoma

doederleini Rowe & Pawson, 1977. A. Reticulated specimen (UF5318). B. Close-up of annulated arm

spines. C. Densely reticulated specimen (IJF6518). D. Ophiocoma brevipes Peters, 1851 (SMNH-

133230). E-F. Ophiocoma dentatayivXlQX & Troschel, 1842. E. Variegated specimen (SMNH-133232).

F. Uniformly reddish brown specimen (URUN2009-11047). G-L. Ophiocoma krohi sp. nov. paratypes.

G. With large patches (SMNH-Type-8533). H. Small irregular spots (MNHN-IE-4302). I. Star-shaped

radiating pattern (SMNH-Type-8531). J. Densely mottled (MNHN-IE-4301). K. Uniformly dark brown

(SMNH-Type-8536). L. Eight brown, sparsely mottled (UF13938). For institution codes see main text.

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

short, externally with two large tooth foramina in dorsal half, dorsalmost foramen oval, wider than long,

other foramen square (Fig. 3H); internally eaeh foramen divided by a wide septum, dorsalmost one into

two small holes, other foramen into two parallel slits (lower septum broken in Fig. 3H2). External ventral

half of dental plate with smooth horizontal ridges, forming two wide ovals, the ventralmost one half the

size of the other; a eluster of tiny depressions at ventral end of plate. Radial plate shaped as a wide triangle,

proximal edge eoneave, distal point with eondyle, depressed at lateral edge (Fig. 3F), but not as ball-like

as in O. brevipes and O. krohi sp. nov. Adradial genital plate long, flat, thin, thiekened at distal end, with

eondyle and pit (Fig. 311); abradial genital plate thin, blade-like, eurved distally (Fig. 312). Oral plates.

Fig. 3. Ophiocoma (Breviturma subgen. nov.) doederleini de Loriol, 1899 (IJF5318). SEM images.

A. Dorsal dise. B. Dorsal arm. C. Dorsal dise granules. D. Jaws. E. Ventral arm. F. Radial shield,

internal aspeet, distal edge downwards. G. Oral plates (half-jaws), abradial aspeet left, adradial right,

proximal edge left. H. Dental plates, external aspeet on left, internal on right (lower septum broken),

dorsal upwards. I. Genital plates abradial left, adradial right, distal upwards. J. Arm vertebrae, proximal

aspeet left, distal right, dorsal upwards. K. Eateral arm plates, external aspeet left, internal right, distal

to the right. UF, Florida Museum of Natural History.

European Journal of Taxonomy 68 : 1-26 ( 2013 )

adradial distal part with strong folds, abradial face with large musele flange with horizontal striations,

2.5 mm wide, 2.0 mm high (Fig. 3G). Pro xim al vertebrae are 2.4 mm wide and 1.8 mm high (Fig. 3J).

UF6512

Collecting data. La Reunion Island, Saint Gilles, Point des Aigrettes, 21°02’11” S, 55°13’01” E, fore

reef, under roeks, 6-22 m dive, eolleetors H. Bruggemann, N. Hubert, F. Miehonneau, G. Paulay, 12

Aug. 2007, 1 spm.

Morphological data. 17 mm dd. Granule density 88-96 mm'^. Arm spine sequenee 3, 3, 3, 4, 4, 5, 5, 5,

5, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, eontinued until the distal arm where the spines deerease to three. Brown

dorsal dise with retieulating net pattern. Dorsal arms dark brown with widely spaeed short white bands

with dark markings. Ventral dise dark and light brown mottled. Oral frame white with few dark spots.

Ventral arms dark brown with few short white bands. Spines white, strongly annulated with brown rings,

long and slender, dorsalmost spine barely two segments long, middle spines longer than two segments.

UF 6634

Collecting data. La Reunion Island, Saint-Leu See Jaune, 21°12’20” S, 55°16’58” E, under roeks, 12-

15 m dive, eolleetors H. Bruggemann, N. Hubert, F. Miehonneau, G. Paulay, 17 Aug 2007, 1 spm.

Morphoeogicae data. 26 mm dd. Granule density 60-64 mm ~^. Arm spine sequenee 3, 3, 3, 4, 4, 5, 5, 5,

5, 5, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4.3. Dorsal dise medium brown with dark mesh pattern (retieulated),

dorsal arms dark brown with widely spaeed lighter brown bands. Oral frame ereamy white with brown

spots and irregular markings, ventral dise light brown with dark retieulations, ventral arm medium

brown with widely spaeed eream markings. Spines white, strongly annulated with brown rings, long

and slender, dorsalmost spine barely two segments long, longest middle spine almost as long as three

segments.

UF6518 (only image available)

CoEEECTiNG DATA. La Reunion Island, Saint Gilles, Point des Aigrettes, 21°02’11” S, 55°13’01” E, fore

reef, under roeks, 6-22 m dive, eolleetors H. Bruggemann, N. Hubert, F. Miehonneau, G. Paulay, 12

Aug. 2007, 1 spm.

Morphoeogicae data. 12.5 mm dd. Colour pattern on dorsal dise eonsists of thieker, brown, retieulating

lines, instead of a thin net-like pattern as in the other speeimens; otherwise similar to the others (Fig. 2C).

Remarks

Ophiocoma doederleini oeeurs in several eolour morphs, with various retieulated patterns (Fig. 2A, C) or

spotted dise (Devaney 1970), whieh is eonflrmed by our moleeular data (Boissin & Hoareau, unpublished).

It ean easily be distinguished from the other speeies of Breviturma subgen. nov. by its dark and light

strongly annulated, more slender, and longer arm spines. It also has the widest dorsal arm plate of all

speeies of Breviturma subgen. nov. The high granule density in the smallest speeimen was surprising and

may need further investigation. Maximum size at least 26 mm aeeording to the studied material.

Ophiocoma {Breviturma) brevipes Peters, 1851

Figs 2D, 4A-E, 5A, D, G, J, M, P

Ophiocoma brevipes VqXqvs, 1851: 466.

Ophiocoma brevispinosa Smith, 1876: 40.

Ophiopeza danbyiTdiXiyJhdiX, 1897: 189.

non Ophiocoma brevipes - Stohr et al. 2008: 553.

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

Diagnosis

Species of Ophiocoma (Breviturma) subgen. nov. characterized by a light colour pattern, cream to white

with some brown to grey markings; five arm spines from segment 4 or 5 on large specimens for more

than 10 segments; disc granule densities up to about 250 mm'^. Maximum size at least 20 mm dd

according to the examined material.

Type material examined

Syntype ZMB Ech 961

Collecting data. Mozambique, 18° 39.942’ S 35° 31.774’ E (from atlas), collector Peters, 1851.

Morphological data. 12-13 mm dd (irregular disc), white dorsal and ventral, any pattern bleached,

hardly visible. DAP W:L 2:1, arm spine sequence 3, 3, 4, 4, 5, 5, 5, 5, 5, 4, 4, 4, 4, 4, 4, oral shield longer

than wide, ventral disc completely granulated, 2^^ ventral arm spine longest, longer than an arm segment.

Syntype ZMB Ech 962

Collecting data. Mozambique, 18° 39.942’ S 35° 31.774’ E (from atlas), collector Peters, 1851.

Morphological data. 13 mm dd, white dorsal and ventral disc, dorsal arms with brown bands, ventral

arms white, DAP W:E 2:1, arm spine sequence 3, 3, 4, 5, 5, 5, 5, 5, 5, 5, 5, 5, 5, 5, 4, 4, 4, oral shield

longer than wide, ventral disc completely granulated, 2"^ ventral arm spine longest, longer than an arm

segment.

Syntype ZMB Ech 963

Collecting data. Mozambique, 18° 39.942’ S 35° 31.774’ E (from atlas), collector Peters, 1851.

Morphological data. 13 mm dd, dors ally whitish-brown colour pattern, ventrally white, DAP W:E =

2:1, arm spine sequence 3, 3, 3, 4, 5, 5, 5, 5, 5, 5, 5, 4, 4, 4, 4, 4, 4, oral shield about as wide as long,

ventral disc completely granulated, 2"^ ventral arm spine longest, longer than an arm segment.

Syntypes ZMB 4660

Collecting data. Mozambique, Querimba Islands, 12° 25.796’ S 40° 35.948’ E (from atlas), collector

Peters, 1851, 3 spms.

Morphological data. Spm A, 17-18 mm dd, white overall, DAP W:E 2:1, arm spine sequence 3, 3, 3,

4, 4, 5, 5, 5, 5, 5, 5, 5, 4, 4, 4, 4, 4, oral shield longer than wide; Spm B, 15 mm dd, white, DAP W:E

2:1, arm spine sequence 3, 3, 3, 4, 4, 5, 5, 5, 5, 5, 5, 4, 4, 4, 4, 4, 4, oral shield wider than long; Spm

C,12 mm dd, white, DAP W:E 2:1, arm spine sequence 3, 3, 3, 4, 4, 5, 5, 5, 5, 5, 5, 5, 5, 5, 5, 5, 5, oral

shield longer than wide; all specimens ventral disc completely granulated, 2"^ ventral arm spine longest,

longer than an arm segment.

Other material examined

SMNH-133230

Collecting data. Ea Reunion, Etang Sale, 21°16’17 S, 55°19’56 E, collectors E. Boissin, T. Hoareau,

24 Feb. 2009. 1 spm.

Morphological data. 13 mm dd, dorsal disc grey-brown with a mosaic of irregular white spots (Fig. 2D),

dorsal arms grey-brown with some dark spots, spines with dark longitudinal line, tips white, ventral disc

white, grey-brown marbled, oral frame white, oral shields with dark brown spots, ventral arms white,

arm spine sequence 3, 3, 3, 4, 4, 5, 5, 5, 5, 5, 4, 4, 4, 4, 4, 4, two dorsal spines shorter than arm width.

European Journal of Taxonomy 68 : 1-26 ( 2013 )

Fig. 4. SEM images of three speeies of Ophiocoma (Breviturma) subgen. nov. A-E. Ophiocoma brevipes

Peters, 1851 (SMNH-133230). A. Dorsal dise. B. Dorsal arms. C. Dorsal dise granules. D. Jaws.

E. Ventral arm. E-I. Ophiocoma dentata Muller & Trosehel, 1842 (SMNH-133232). E. Dorsal dise.

G. Dorsal arm. H. Dorsal dise granules. I. Jaws. J. Ventral arm. K-N. Ophiocoma krohi sp. nov. paratype

(SMNH-Type-8534). K. Dorsal dise. L. Dorsal arms. M. Dorsal dise granules. N. Jaws. O. Ventral arm.

For institution eodes see main text.

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

2^^ (and third) ventral spine longest. Ventral interradii fully granulated. Granule density on light image

248-280 mm ~^.

Dorsal dise granule diameter 42-50 pm, granule density 240-248 mm'^, on SEM image (Fig. 4A, C);

third spine about 1 mm long, proximal DAPs about 1.5 mm wide (Fig. 4B). Oral shield as wide as

long; 6-8 tooth papillae, 5 lateral oral papillae at eaeh jaw edge, outer papilla widest (Fig. 4D). Few

granules on distal ends of oral shields. Two tentaele seales (Fig. 4E). Oral plates, adradial distal part with

strong folds, abradial faee with large musele flange with horizontal striations, 2.4 mm wide, 2.0 mm

high (Fig. 5A). Dental plate short, externally with two large tooth foramina in dorsal half, dorsalmost

foramen oval, wider than long, other foramen almost round, internally eaeh foramen divided by a wide

septum, dorsalmost one into 2 long parallel slits, other one to 2 short oval holes; on external ventral

half of dental plate two raised horizontal ridges, at edge a eluster of tiny depressions (Fig. 5D). Radial

shield angular, internally with distal large round eondyle (Fig. 5G). Adradial genital plate long, flat,

thin, thiekened at distal end, with eondyle and pit; abradial genital plate thin, blade-like, eurved distally

(Fig. 5J). Proximal vertebrae 2.0 mm wide and 1.7 mm high (Fig. 5M).

URUN 2013-11188

Collecting DATA. Fa Reunion, Saint Pierre, 21°20’27 S, 55°27’31 E. 1 spm.

Morphological data. 19.7 mm dd. Arm spine sequenee 3, 3, 3, 4, 4, 5, 5, 5, 5, 5, 5, 5, 5, 5, 5, 5, 5, 5, 4,

4, 4, 4, 4, 4. Granule density 176-188 mm'^.

URUN 2008-04774

Collecting data. Madagascar, station MGNW-9, Sakatia Island, 13°18’16 S 48°08’51 E. 1 spm.

Morphological data. 13.08 mm dd. White with brown marbled pattern on dorsal and ventral dise,

dorsal arms brown, oral frame and ventral arm white, few dark spots on oral and adoral shields. Granule

density 148-168 mm'^. Arm spine sequenee 3, 3, 3, 4, 4, 5, 5, 5, 5, 5, 4, 4, 4, 4, 4, 4.

URUN 2008-05819

Collecting data. Madagascar, station MGNW-9, Sakatia Island, 13°18’16 S, 48°08’51 E. 1 spm.

Morphological data. 13.5 mm dd. White-brown to grey marbled disc, grey arms with faint dark median

line, oral frame and ventral arms white, spines mostly white. Granule density 180-200 mm ~^. Arm spine

sequence 3, 3, 3, 4, 4, 5, 5, 5, 5, 5, 5, 4, 4, 4, 4, 4, 4, 4.

Remarks

Ophiocoma brevipes is clearly distinguished from the other species of Breviturma subgen. nov. by its

consistently white and light brown to grey colour pattern, up to five arm spines far out on the arms, and

dorsal disc granule densities up to 250 mm'^. According to Devaney (1970), live spines occur in some

specimens Ifom 3 mm dd.

Ophiocoma {Breviturma) dentata Muller & Troschel, 1842

Figs 2E, F, 4F-J, 5B, E, H, K, N, Q

Ophiocoma dentata Muller & Troschel, 1842: 99, pi. 7, flgs 3-3a.

Ophiocoma ternispina v. Martens, 1870: 252.

Ophiocoma insularia Lyman, 1862: 80.

Ophiocoma variegata Smith, 1876: 39.

non Ophiocoma marmorata - Devaney 1970: 17.

European Journal of Taxonomy 68 : 1-26 ( 2013 )

Diagnosis

Species of Ophiocoma (Breviturma) subgen. nov. characterized by rarely possessing more than four

arm spines and if present only at few segments; arm spines never annulated; variable in colour pattern

(uniformly or marbled brown, reticulated of spotted); dorsal disc granule density less than 150 mm'^.

Maximum size at least 28 mm dd (Devaney 1970).

Type material examined

Holotype ZMB Ech 931

Holotype of Ophiocoma dentata.

Collecting data. Celebes (Sulawesi), 1° 50.875’ S 120° 31.675’ E (from atlas), collector Deppe.

Morphological data. 18-19 mm dd (irregular disc), granules on dorsal disc mostly rubbed off, light and

dark brown pattern on dorsal disc, dorsal arms medium brown, ventral interradii fully granulated, white,

DAP W:L 2:1, arm spine sequence 3, 3, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 3, 3, 3, 3, 3, oral shield longer than

wide, ventralmost and 2"^^ ventral spine longest, longer than an arm segment, dorsal spines thickened,

ventral ones slender, not annulated.

Holotype ZMB Ech 1815

Holotype of Ophiocoma ternispina.

Collecting data. Indonesia, Flores-Sea, Larantuka, Flores, 8° 21.4’ N 122° 57.817’ E (from atlas),

1 spm.

Morphoeogicae data. 16 mm dd, disc light brown, triangular brown spots and curved lines on DAPs,

spines weakly annulated, DAP W:F 2:1, disc irregular round, ventral interradii completely granulated,

arm spine sequence 3, 3, 3, (3)4, 4, 4, (4)3, 3, 3, 3, 3, 3, 3, 3, 3,3, 3 (numbers vary between arms), 2"^

ventral spine longest, slender spines, oral shield twice as long as wide.

Syntype SMNH-Type-5244

Syntype of Ophiocoma insularia.

CoEEECTiNG DATA. Hawaii, 1860, 1 spm (originally donated to Stockholm by Museum of Comparative

Zoology at Harvard).

Morphoeogicae data. 17.6 mm dd, disc light brown marbled, arms light brown, dorsal and ventral

side similar in colour, disc fully granulated dorsally and ventrally. No annulations on arm spines, spine

sequence 3, 3, 3, 4, 4, 4, 5(4), 5(4), 5(4), 4, 4, 4, 4, 4, 4, 4, 4, 4, 3, 3, 3 (two arms with only 4 spines).

Oral shield oval. Granule density 96-120 mm'^.

Syntypes MCZN 1668

Syntypes of Ophiocoma insularia.

CoEEECTiNG DATA. Hawaii, Maui, 1860, 7 spms.

Morphoeogicae data. 13.3-16.6 mm dd, all dark brown to black on dorsal disc and arms, including

spines, ventral side slightly lighter brown. Both middle spines about equal, longer than dorsal and

ventral spines. (Photos provided by Adam Baldinger, MCZN).

Syntypes MCZN 1666

Syntypes of Ophiocoma insularia.

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

Fig. 5. SEM images of skeletal elements of Ophiocoma (Breviturma subgen. nov.) brevipes Peters,

1851 (SMNH-133230), O. (B.) dentata Muller & Trosehel, 1842 (SMNH-133232) and O. krohi sp. nov.

paratype (SMNH-Type-8535). A-C. Oral plates, abradial aspeet at left, orientated with dorsal side up. A.

O. brevipes. B. O. dentata. C. O. krohi sp. nov. D-F. Dental plates, external aspeet left, internal aspeet

right, dorsal edge upwards. D. O. brevipes. E. O. dentata. F. O. krohi sp. nov. G-I. Radial shields, internal

aspeet, proximal edge upwards. G. O. brevipes. H. O. dentata. I. O. krohi sp. nov. J-L.Vertebrae from

proximal part of arm, proximal aspeet left, dorsal upwards. J. O. brevipes. K. O. dentata. L. O. krohi

sp. nov. M-O. Genital plates, abradial plate left, adradial plate right, distal ends up (towards radial

shields). M. O. brevipes. N. O. dentata. O. O. krohi sp. nov. P-R. Lateral arm plates from proximal

arm, external aspeet left, internal aspeet right, distal end right. P. O. brevipes. Q. O. dentata. R. O. krohi

sp. nov. For institution eodes see main text.

European Journal of Taxonomy 68: 1-26 (2013)

Collecting data. Hawaii, 1860, 2 spms.

Morphological data. 19.5 and 31.6 mm dd, dark brown disc and arms, with lighter abraded patehes on

dorsal dise, dark brown on ventral dise and arms; a note among the labels reads “4-5 spines, 3 upper

about equal”. (Photos provided by Adam Baldinger, MCZN).

Other material examined

SMNH-133232

Collecting data. La Reunion, Etang Sale, 21°16’17” S 55°19’56” E, eolleetors E. Boissin, T. Hoareau,

24 Feb. 2009, 1 spm.

Morphoeogicae data. 11 mm dd, dorsal dise with a brown multipointed star-shape, white eentre spot with

brown dots, interradii white with brown marbled pattern (Fig. 2E); dorsal arm white with reetangular dark

brown outlines, spines brown at base, fading into white towards their tip. Ventral interradii brown and

white marbled, fully granulated to oral shields, oral frame white, oral shields with 2 parallel longitudinal

brown lines. Ventral arm white with median brown dots and short lines. Arm spine sequenee 3, 3, 3, 3,

4, 4, 4, 4, 4, 4, 3, 3, 3, 3, 3, 3. Granule density 216-224 mm'^.

Dorsal dise granule diameter 49-54 pm, granule density 120-144 mm'^, on SEM image (Fig. 4F, H);

third spine on proximal segments 1 mm long, DAPs 1.4-1.6 mm wide Fig. 4G). Oral shields as wide

as long; 5-6 tooth papillae, 5 lateral oral papillae at eaeh jaw edge, outer papilla widest (Fig. 41). Few

granules on proximal ends of oral plates. Oral plates, adradial distal part with strong folds, abradial

faee with large musele flange with horizontal striations (Fig. 5B), 1.6 mm wide, 2.0 mm high. Dental

plate short, externally with two large tooth foramina in dorsal half, dorsalmost foramen oval, wider than

long, other foramen square; internally eaeh foramen divided by a wide septum, dorsalmost one into two

wide vertieal oval slits (septum broken), other foramen into two short parallel holes (Fig. 5E). External

ventral half of dental plate with smooth horizontal ridges, forming two wide ovals, the ventralmost one

half the size of the other; a eluster of tiny depressions at ventral end of plate (Fig. 5E1). Radial shield

triangular with eoneave proximal edge, internally with distal round eondyle (Fig. 5H), not as ball-like

pronouneed as in O. brevipes and O. krohi sp. nov. Adradial genital plate (Fig. 5K1) long, flat, thin,

thiekened at distal end, with eondyle and pit; abradial genital plate (Fig. 5K2) thin, blade-like, eurved

distally. Proximal vertebrae 1.6 mm wide and 1.2 mm high (Fig. 5N).

URUN 2009-11047

CoEEECTiNG DATA. Fa Rcunion, Etang Sale, 21°16’17” S 55°19’56” E, eolleetors E. Boissin, T. Hoareau,

24 Feb. 2009, 1 spm.

Morphoeogicae data. 23.7 mm dd, dorsal dise uniformly reddish brown (Fig. 2F), dorsal arm dark

brown, ventral dise and arms dark brown. Granule density 44-60 mm'^, granule diameter 70-90 pm. Arm

spine sequenee 3, 3, 3, 4, 4, 4, 4 (5), 4 (5), 4, 4, 4, 4, 4, 3, 3, 3, 3 (5 spines on one side of one arm, onee

at segment 7 and onee at segment 8).

URUN 2007-00380

CoEEECTiNG DATA. Fa Reunion, Fa Saline Trou d’eau, 21°06’09” S 55°14’24” E, eolleetor G. Paulay,

2007, 1 spm.

Morphoeogicae data. 24.2 mm dd, dorsal dise brown, dorsal arms dark brown with widely spaeed light

markings, spine slighter brown, ventral dise brown, oral frame lighter, ventral arms brown. Granule

density 52-68 mm'^. Arm spine sequenee 3, 3, 3, 4, 4, 4, 4, 4, 4, 4, 4, 4, 3, 3, 3, 3, 3.

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

Remarks

The types of O. insularia are elearly all eonspeeifie. Their eolouration and large size suggest that they are

also eonspeeifie with O. dentata. Ophiocoma dentata oeeurs in several eolour morphs, from uniformly

dark brown to brown marbled dises with banded arms, and retieulated or spotted forms that resemble

O. doederleini. It ean always be distinguished from the latter speeies by the absenee of annulations

on the arm spines, and shorter arm spines. In addition, only the largest individuals sometimes have

more than four spines on few arm segments. The eonspeeifieity of the various eolour morphs is also

eonfirmed by moleeular data (Boissin & Hoareau, unpublished). Another speeies eurrently believed to

be eonspeeifie with O. dentata is O. variegata with type loeality Rodriguez Island. Aeeording to the

original deseription (Smith 1876), the type measured 28 mm dd and had only four arm spines, whieh

agrees with O. dentata.

Devaney (1970) synonymized O. marmorata with O. dentata, but as we diseuss below, the holotype of

O. marmorata is most likely not eonspeeifie with O. dentata.

Ophiocoma (Breviturma) krohi sp. nov.

um:lsid:zoobank.org:aet:D662D2A3-33QD-4693-8757-E6112C4BABCl

Figs 2G-L, 4K-0, 5C, F, I, F, O, R, 6

Ophiocoma brevipes - Stohr et al. 2008: 553.

Ophiocoma sp. nov. - Hoareau et al. 2013: 2.

Diagnosis

Speeies of Ophiocoma {Breviturma) subgen. nov. with variegated to uniform dark brown eolour pattern;

arm spines may be weakly annulated, but not as elearly light and dark banded as in O. doederleini', up to

four arm spines (in large speeimens beyond segment 15), sometimes five at segments 6-9 (rarely 5-8);

relatively eoarse dise granulation. Maximum size about 16 mm dd.

Etymology

The speeies is named in honour of Dr. Andreas Kroh, Vienna, for his important eontributions to

eehinoderm systematies.

Type material

Holotype

Fa Reunion Island, St. Pierre, 21°20’27 S 55°27’31 E, eolleetors E. Boissin, T. Hoareau, 25 Mar. 2009,

in ethanol (MNHN-IE-4300).

Paratypes

From same loeality as holotype 8 spms: 1 spm, disseeted, fragments on SEM stubs and arms in ethanol

(SMNH-Type-8535); 1 spm gold eoated anduneoated arms dry (SMNH-Type-8534); 2 spms in ethanol

(SMNH-Type-8531, 8536); 1 spm in ethanol (UF13938); 1 spm in ethanol (URUN 2009-11108); 2 spms

in ethanol (MNHN-IE-4301, 4302). Fa Reunion, Cap Homard, 21°02’07” S 55°13’06” E, eolleetors

E. Boissin, T. Hoareau, 23 Feb. 2009: 2 spms in ethanol (SMNH-Type-8532, 8533); 1 spm in ethanol

(MNHN-IE-4303).

Description

Holotype

12.7 mm dd, radial shields eompletely obseured by granules, granule density 104 mm'^. Arm spine

sequenee 3(2), 3(2), 3, 4, 4, 4, 4, 4, 4, 4, 4, 3, 3, 3, 3. Dorsal dise with dark and lighter brown lines

European Journal of Taxonomy 68: 1-26 (2013)

radiating from a dark central patch, some white spots at disc edge (Fig. 6A). Dorsal arms medium

brown with dark and white transverse bands (Fig. 6B). Ventral disc dark brown mottled, ventral arms

and oral frame lighter brown, oral shields dark. Ventral interradii completely, but sparsely granulated

(Fig. 6D, E). Dorsal spines brown, ventral spines weakly annulated brown on white. DAPs oval, W:L

1.8:1, contiguous. Dorsalmost spine a little longer than a segment, middle spines about 2 segments long,

ventralmost spine slightly shorter. Arm spines flattened, slightly tapered with truncated tips, dorsalmost

spine wider than other spines. Oral shields trapezoid, distally almost as wide as long, proximally just

over half as wide as long, lateral edges straight (except madreporite), proximal and distal edges convex

(Fig. 6D). Madreporite larger than other oral shields, more rounded, all edges convex. At tip of jaw a

cluster of 8-10 small, round tooth papillae. Along each jaw edge 4 rectangular, wider than high, oral

papillae, and distally a smaller papilla at an angle to the others (Fig. 6E). Adoral shields uneven triangular,

framing the lateral edges of the oral shield, not meeting proximally. Bursal slits extend from oral shield

to disc edge. First ventral arm plate outside mouth-slit, small, winglike pentagonal, middle of distal

edge notched. Following plates pentagonal, about as long as greatest width, with obtuse proximal angle,

lateral edges notched, distal edge convex, contiguous (Fig. 6C). 2 oval scales at a slightly open angle at

the lateral arm plate of each tentacle pore proximally; the inner scale decreases gradually and is absent

on the distalmost segments. Tube feet expanded, smooth, with distal constrictions and terminal bulb.

The paratypes are similar to the holotype in most characters, except as follows:

SMNH-Type-8534

1 spm, 11 mm dd, dark brown radiating stripes on dorsal disc, interradii mottled light and dark brown,

overall small white dots; dorsal arms dark brown with some lighter markings, spines white with diffuse

brown pattern, some with annulations (not flgured). Ventral interradii dark brown mottled with white,

fully granulated; adoral shields and oral plates unevenly brown, oral shields dark brown with white

edges, oral papillae white. Ventral arm plates and tentacle scales brown with white edges. Arm spine

sequence 3, 3, 3, 4, 4, 5, 5, 4, 4, 4, 4, 4, 4, 4, 4, 4. Granule density 132-136 mm'^ on light image.

Fig. 6. Holotype of Ophiocoma (Breviturma subgen. nov.) krohi sp. nov. (MNHN-lE-4300). A. Dorsal

disc. B. Dorsal arm. C. Ventral arm, note the pattern on the arm spines. D. Ventral disc. E. Jaws.

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

Dorsal disc granule diameter 57-73 pm, granule density 132-136 mm'^, on SEM image (Fig. 4K, M);

third arm spine 1.3 mm long, DAP 1.2 mm wide (Fig. 4F). Oral shields as wide as long (Fig. 4N). 5

tooth papillae, 5 lateral oral papillae at each jaw edge, outer papilla widest. Few granules on proximal

ends of oral plates (Fig. 4N).

SMNH-Type-8535

1 spm, 13 mm dd, dorsal disc dark brown, white spots on radial shields; dorsal arms banded dark and

light brown; spines dark brown, on distal arm annulated (not figured). Granule diameter 57-74 pm,

density 104 mm'^. Ventral interradii dark brown; adoral shields light brown with dark spots; oral shields

dark brown with white edges; oral papillae white. Ventral arm plates brown with white edges and lighter

areas. Arm spine sequence 3, 3, 3, 4, 4, 5, 5, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4. Up to 5 tooth papillae, 5 lateral oral

papillae at each jaw edge, outer papilla widest. Few granules on distal ends of oral plates.

Oral plates, adradial distal part with strong folds, abradial face with large muscle fiange with horizontal

striations that end in deep incisions in the distal edge; 1.9 mm wide, 2.3 mm high (Fig. 5C). Dental

plate short, with two tooth foramina on external dorsal half, dorsalmost one oval, wider than long,

other small, round; internally a wide septum divides each foramen into two pairs of vertical oval holes,

upper ones larger than lower ones (Fig. 5F). Radial shield angular, with large round condyle at internal

distal end (Fig. 51). Adradial genital plate long, fiat, thin, thickened at distal end, with deep pit; abradial

genital plate thin, blade-like, curved distally (Fig 5F). Proximal vertebrae 2.0 mm wide and 1.5 mm high

(Fig. 5Q).

SMNH-Type-8531

1 spm, 6.8 mm dd, granule density 152 mm'^. Arm spine sequence 3, 3, 3, 4, 4, 5, 5, 4, 4, 4, 4, 3, 3,

3, 3, 3, 3. Dorsal disc with central dark brown patch, with white central spot, dark brown radiating

lines, medium brown interradial lines on light brown background, white spots at disc edge (Fig. 21).

Dorsal arms with transverse dark and lighter brown bands. Ventral disc light brown with irregular cream

patches, oral frame and ventral arms lighter brown, oral shields dark.

SMNH-Type-8532

1 spm, 9.6 mm dd, granule density 116-120 mm'^. Arm spine sequence 3, 3, 3, 4, 4, 4, 4, 4, 4, 4, 4, 4, 3,

3,3,3. Dorsal disc dark brown mottled, dorsal arms lighter brown with dark bands. Ventral disc light and

dark brown mottled (not figured). Ventral arms and oral frame light brown, oral shields with dark patch.

MNHN-IE-4303

1 spm, 8.1 mm dd, granule density 124 mm'^. Arm spine sequence 3, 3, 3, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 3,

3, 3, 3. Dorsal disc light brown mottled, dorsal arms light brown with dark bands. Ventral disc light and

dark brown mottled, arms and oral frame creme, oral shields with dark patches.

SMNH-Type-8533

1 spm, 10 mm dd, granule density 140-164 mm'^. Arm spine sequence 3, 3, 3, 4, 4, 5, 5, 5, 5, 4, 4, 4, 4,

4, 4, 4, 4, 4, 4, 4,4, 3, 3, 3. Dorsal disc variegated white and brown, arms dark and lighter brown banded,

spines annulated (Fig. 2G).

SMNH-Type-8536

1 spm, 15.5 mm dd, granule density 64-96 mm'^. Arm spine sequence 3, 3, 3, 4, 4, 4, 4, 4, 4, 4, 4, 4,

4, 4, 4, 4, 4, 4, 4, 4, 3, 3, 3. Dorsal disc uniformly dark brown, except on radial shields white granules

(Fig. 2K). DAPs dark brown, all spines weakly annulated with irregular dark stripes on white. Oral

frame white with dark marks on oral shields. Ventral arm plates dark brown with white borders.

European Journal of Taxonomy 68: 1-26 (2013)

MNHN-IE-4302

1 spm, 13 mm dd, granule density 96-104 mm'^. Arm spine sequenee 3, 3, 3, 4, 4, 4, 4, 4, 4, 3, 3, 3, 3,

3, 3. Light brown dise with irregular dark spots or blotehes. Dorsal arms medium brown with widely

spaeed dark brown transverse bands (Fig. 2H). Ventral dise medium and dark brown mottled, ventral

arms and oral frame light brown to eream, oral shields with dark pattern.

MNHN-IE-4301

1 spm, 11.8 mm dd, granule density 72-96 mm'^. Arm spine sequenee 3, 3, 3, 4, 5, 5, 5, 5(4), 4,4(5), 4, 4,

4, 4, 4, 4, 4, 4, 4, 3, 3, 3. Dorsal dise dark brown mottled, dorsal arms dark brown with few white marks,

spines dark brown (Fig. 2J). Oral frame eream white with light brown marks on oral shields, ventral arm

plates brown.

URUN 2009-11108

1 spm, 11.3 mm dd, granule density 112-120 mm'^. Arm spine sequenee 3, 3, 3, 4, 4, 4, 4, 4, 4, 4, 4, 4,

4, 3, 3, 3, 3, 3, 3, 3, 3. Dorsal dise with dark brown star-shaped pattern, interradii brown with irregular

white spots, irregular white spots in eentre of dise, white granules on radial shields (not figured). Dorsal

arms mostly dark brown, some lighter patehes, spines dark brown. Oral frame white, brown patehes on

oral shields. Ventral arm plates brown with white margins.

UE13938

1 spm, 10.3 mm dd, granule density 112-116 mm'^. Arm spine sequenee 3, 3, 3, 4, 4, 5, 5, 5, 5(4), 4, 4, 4,

4, 4, 3, 3, 3. Dorsal dise light brown with irregular dark patehes and spots, dorsal arm light brown with

widely spaeed narrow dark brown bands (Fig. 2L). Spines light brown, weakly annulated. Oral frame

white with light brown pattern on oral shields, ventral arm plates light brown with white margins.

Remarks

Ophiocoma krohi sp. nov. differs from O. brevipes and O. dentata in maximum size, granule size

and density, and in eolour pattern. Although quite variable in eolour, it has never been observed with

retieulated or spotted pattern as O. dentata and O. doederleini. We examined 12 speeimens and half of

them showed up to five arm spines on 2-4 segments, the other half had only four. Thus they all differ in

spine number from O. brevipes, but some are similar to O. dentata in laeking a fifth spine, although they

may have four spines farther out on the arm than O. dentata.

The speeimens reported by Stohr et al. (2008) are most likely eonspeeifie with O. krohi sp. nov. based

on eolour pattern, the presenee of five spines and their small size. The speeies has a wide geographie

distribution on eoral reefs aeross the Indian and Paeifie Oeeans at depths of 0-30 m (Hoareau et al.

2013).

non-Breviturma, unelear subgenus

Ophiocoma marmorata Marktanner-Tumeretseher, 1887

Op/z/ocowa warworato Marktanner-Tumeretseher, 1887: 303, pi. 12 figs 16, 17.

non Ophiocoma dentata - Devaney 1970: 17.

Material examined

NHMW 3. Zool. Abt. Nr. 10.466

Colleetion Eiehhom 2873, 0-7°N, 23-25°W (= Tropieal Atlantie). Lot of 5 spms, wrongly registered

as “syntypes”. One very obviously belongs to another speeies and was not examined more elosely.

Examined, photographed and data transmitted by A. Kroh, Vienna. Eaeh spm is treated separately below.

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

Description

Holotype NHMW 10.466a

8.7 mm dd, dorsally light orange eolour pattern with darker spots on arms, ventrally eream eoloured,

spines white, not annulated, arm spine sequenee (2)3, 3, 3, 3(4), 4, 4, 4, 3(4), 4, 3(4), 4(3), 3(4), 4(3),

3, 4, 3 (numbers vary between arms and alternate between segments), oral shield longer than wide.

Granule density about 80 mm'^. Holotype status inferred from original deseription.

NHMW 10.466b

6.3 mm dd, eream eolour dorsal and ventral, no pattern, eoarser granules than holotype, arm spine

sequenee 3, 3, 3(4), 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4. Granule density 60 mm'^.

NHMW 10466c

6.1 mm dd, eream eolour dorsal and ventral, no pattern, eoarser granules than holotype, arm spine

sequenee 3, 3, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4, 4. Granule density 48 mm'^.

NHMW 10.466d

5.2 mm dd, eream eolour dorsal and ventral, no pattern, eoarser granules than holotype, arm spine

sequenee 3, 3, 4, 4, 4, 4(5), 4(5), 4, 4, 4, 4, 4, 4, 4, 4. Granule density 60 mm'^.

Remarks

The original deseription of O. marmorata mentions only a single speeimen of 9 mm dd (Marktanner-

Tumeretseher 1887), whieh should be regarded as the holotype. The remaining three speeimens eannot be

regarded as type material sinee they are not mentioned in the deseription. The loeality of O. marmorata is

most likely ineorreet as it lies in the middle of the Atlantie Oeean at great depth. Ophiocoma is a shallow

water tropieal genus, rarely reported below 100 m depth. Taking into aeeount that the determination of

eoordinates, in partieular longitude, was diffieult and often inaeeurate in the 19^*^ eentury, we think it

possible that the eorreet loeality may be the Saint Peter and Saint Paul Arehipelago, whieh is situated at

0.9°N, 29°W, assuming that the data are based on the Greenwieh Meridian. Instead, the Ferro Meridian

may have been used, whieh would put the loeality at 40°40’-42°40’W today, even eloser to Brazil.

The speeimens were bought by direetor Eiehhom of Graz (Austria) from the trader Hugo Sehilling

in Hamburg in 1882, together with several other eehinoderms, and transferred to the natural history

museum in Vienna (A. Kroh, pers. eomm.).

The alternating spine sequenee of the holotype of O. marmorata may be interpreted as an unusual

variation, if we aeeept that it is eonspeeifie with the three smaller speeimens in the lot. This eharaeter is

typieal for speeies in the scolopendrina group though and therefore the holotype of O. marmorata may

belong to one of the speeies in that group, possibly O. echinata (Lamarek, 1816) or O. wendtii Muller &

Trosehel, 1842, whieh are the only two speeies of that group known from the Atlantie Oeean. The smaller

speeimens, however, may belong to a different speeies, possibly O. pumila Ltitken, 1856, a member of

the pumila group, whieh has a spine sequenee similar to the smaller O. marmorata (NHMW 10.466b-d)

and is known from the Atlantie. Devaney (1970) does not seem to have examined O. marmorata and did

not know about the existenee of five spines on one of the speeimens in the lot. He also did not eonsider

the signifieanee of geographie distribution when he suggested that O. marmorata (an Atlantie speeies)

may be eonspeeifie with O. dentata (an Indo-Paeifie speeies). Although the fifth arm spines in the small

O. marmorata are in exaetly the same position, arm segments 6 and 7, as in O. krohi sp. nov., the granules

in these small speeimens are high and pointed, different from those found in the speeies of Breviturma

subgen. nov. Granules like that are known from the pumila group. The holotype of O. marmorata has

low round granules, whieh also supports that it is a different speeies from the three smaller speeimens.

European Journal of Taxonomy 68: 1-26 (2013)

The taxonomic status of O. marmorata could not be completely resolved by this study and requires

further investigation, but we strongly doubt that it is conspecific with O. dentata. We propose that the

synonymy of O. dentata includes only O. ternispina, O. insularia and O. variegata.

Remarks on species delimitations

Prior to Devaney’s (1970) work, the three species previously recognized in the brevipes group were

considered mere variations of a single species. Devaney (1970) was able to separate them by spine

sequence, colour pattern and arm width. In addition to the annulated colour pattern of the arm spines,

spine numbers seem to be a critical character to distinguish between O. dentata and O. doederleini.

According to Devaney (1970), O. dentata has five arm spines only at disc diameters above 20 mm and

spine numbers decrease to three on the middle arm (beyond segment 17 at 13-14 mm dd, beyond segment

25 at 24 mm dd), whereas four spines are found on far more distal arm segments in O. doederleini of

similar disc sizes (to segment 32-38 at 13-14 mm dd, to segment 55 at 24 mm dd). In light of our findings

(Table 1) we cannot exclude though that the specimens regarded as O. dentata by Devaney actually

included individuals of O. krohi sp. nov.. All of his data must therefore be treated with caution. The type

of O. dentata (18-19 mm dd) has up to four spines on the proximal arm segments, decreasing to three

from segment 13, which contradicts Devaney’s (1970) observations. Our largest specimen of O. dentata

has two occurrences of five spines among all ten sides of the arms, and three spines from segment 16.

In the type material of O. brevipes, spine numbers appear to be uncorrelated with disc size; five spines are

present from the fourth, fifth or sixth segment and for a variable number of segments before the number

drops to four and finally three again on the distalmost part of the arm. The smallest type, at 12 mm dd,

shows 12 segments with five spines, whereas the largest, at 17-18 mm dd shows only six segments with

five spines; in both animals there are five spines from the sixth segment. Similarly, spine numbers vary

between specimens on the types of the three nominal species currently regarded as conspecific with

(/}

0 5 10 15 20 25 30

Disc diameter

A o. brevipes

X O. dentata

• O. marmorata

+ O. krohi

" O. insularia

~ O. doederleini

Fig. 7. Granule densities in relation to disc diameter of the examined nominal species of Ophiocoma,

from Table 1.

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

O. dentata. In O. krohi sp. nov., the smallest speeimen has five spines on two segments, while the largest

one has only up to four spines. These variations are eommon in the genus Ophiocoma as was observed

already by Devaney (1970). We also found signifieant differenees in granule density among speeies

as shown in Fig. 7 (Kruskal-Wallis test, K=8.9055, d.f=3, P =0.0306). Likewise, granule sizes and/or

densities may be subjeet to growth ehanges as suggested by the inverse eorrelation observed between

the granule density and the dise diameter in O. krohi sp. nov. (Spearman test, R=-0.8012, P=0.0006).

The smallest O. krohi sp. nov. had 152 granules/mm^, whereas the largest one had only 64-96 granules/

mm^. Small O. dentata have three times higher granule densities as large ones, whieh suggests that

granule numbers do not inerease with growth. Somewhat eontradietory are the data for O. brevipes,

where speeimens from La Reunion have greater granule densities than similar size speeimens from

Madagasear.

We deeided to examine a rarely used eharaeter, granule density, that has oeeasionally been mentioned in

speeies deseriptions and revisions, but never been used in a systematie way to differentiate Ophiocoma

speeies. Devaney (1970) used granule densities as aeharaeter in his key to the speeies ofthe scolopendrina

group, but only for few of the speeies. Granule densities were eompared between light images and SEM

images of the same animal and the small observed differenees ean probably be attributed to uneven

distribution of the granules on the dise, rather than to a real differenee in aeeuraey between these

methods. Our method of eounting granules on digital images resulted in eonsiderably higher numbers

than what has been published before. However, published values are not always related to size and as we

see in our results, the largest speeimens have larger and fewer granules. It is also possible that eounts on

images are more aeeurate than eounts under the disseeting mieroseope. In any ease, sinee we used the

same method for all speeimens, the differenees between them are valid and important.

Devaney (1970) examined arm span on the lO^*’ free segment, “eomposed of the length of the longest

arm spines on eaeh side of a segment, the breadth of the dorsal arm plate and the breadth of the lateral

arm plates” (eited from Devaney 1970), and found larger arm spans in O. doederleini than in O. dentata.

However, his values overlap and vary uneorrelated to size, and we eonsider this a weak eharaeter that is

diffieult to assess.

Discussion

Phylogeny

The moleeular phylogeny suggests monophyly of the brevipes group with respeet to the scolopendrina

group, whieh supports the establishment of Breviturma subgen. nov. This is the first time that the

moleeular basis of the speeies groups of Ophiocoma deseribed by Devaney (1970) ean be demonstrated.

The phylogeny also revealed four deep lineages within Breviturma subgen. nov. eorresponding to the

three known speeies and the new speeies deseribed here. Within this subgenus, O. doederleini is the most

distantly related, and the new speeies O. krohi seems to be sister taxon to the O. brevipes!O. dentata

elade, even though the data are not strongly supported.

Morphology

Devaney (1970) showed that the dental plate ean be used to divide the genus Ophiocoma into groups.

All our speeimens eoneur with his deseription of the dental plate in the brevipes group, about twiee

as long as wide with small part for tooth granules, and all tooth foramina divided by a wide septum.

The relationships within Breviturma subgen. nov. are diffieult to resolve with morphologieal means.

Benavides-Serrato & O’Hara (2008) showed that the dental plate in the scolopendrina group differs

between speeies, but that does not seem to hold true within Breviturma subgen. nov. There appear to be

slight differenees in the geometry of the oral plates, but it is eurrently unelear if these are signifieant. The

overall shape of the oral plate is the same in all speeies, but it seems to be lower and shorter in O. dentata

European Journal of Taxonomy 68: 1-26 (2013)

than in O. brevipes and O. krohi sp. nov. Similarly, the geometry of the vertebrae seems to differ slightly,

with O. dentata having the smallest vertebra. However, these were not taken from exaetly the same arm

segment and size differenees may eorrelate with ontogenetie age and overall size of the animal.

As with the moleeular data, Ophiocoma doederleini is elearly distinguished from the other three speeies

by morphology. The most eonspieuous differenee is the presenee of annulated arm spines. The spines

are also longer and thinner than in the other speeies. Granule density of the smallest speeimen eoneurs

with both O. dentata and O. brevipes, while the larger speeimens have densities similar to O. dentata. It

is most similar in arm spine numbers to O. krohi sp. nov. whieh also exhibits weakly annulated spines.

However, the new speeies reaehes a mueh smaller size (16 mm dd vs. 26 mm dd) and differs in dise

eolour patterns (Fig. 2G-L).

Ophiocoma brevipes and O. krohi sp. nov. are similar in having up to five arm spines at small sizes, but

differ in the number of segments that have five spines; they are similar also in the shape of the radial shield

with a large ball-like artieulation eondyle that is less pronouneed in O. dentata and O. doederleini. The

genital plates of the three most elosely related speeies O. brevipes, O. dentata and O. krohi sp. nov. are

similar, but in O. dentata the adradial genital plate is relatively shorter and the abradial genital plate has a

wider proximal edge, similar to O. doederleini. In granule density, O. dentata is elose to O. brevipes, but

both differ greatly from O. krohi sp. nov., whieh again mirrors the assumed phylogenetie relationships.

Conclusion

All four speeies in Breviturma subgen. nov. are similar to eaeh other in some eharaeters. In granule

density and in the presenee/absenee of annulations on the arm spines, O. krohi sp. nov. seems to be

related to O. doederleini. The higher number of arm spines separates O. brevipes from the other three

speeies. Ophiocoma dentata and O. doederleini share several eolour morphs. Ophiocoma krohi sp. nov.

is the smallest of these speeies with about 16 mm dd, a diffieult eharaeter that only elearly distinguishes

signifieantly larger speeimens. It is eurrently impossible to deduee the aneient states of any eharaeters

or infer a phytogeny within the subgenus by morphology. However, a eomprehensive phylogenetie

analysis of the whole genus may offer better elues to solving this problem.

The identity of the new speeies was diffieult to reveal, sinee several nominal speeies have been

synonymized with O. dentata and eaeh one eould potentially be eonspeeifie with the new speeies. The

spine number and granule density appear to be the most important eharaeters to differentiate these

speeies. However, the ophiuroid skeleton grows eontinuously and the number, shape and proportions of

various parts are known to ehange during growth (Stohr 2005). For instanee, we show that in Ophiocoma

krohi sp. nov. the granule density deereases with size. This eomplieates eomparisons with type material

that is naturally only available at limited size ranges and usually not available for disseetion.

The reeent deseription of two new speeies (Benavides-Serrato & O’Hara 2008 and this study) in a

relatively easily aeeessible (shallow water), eonspieuous and well-studied group (ophioeomid brittle

stars) suggests that a eonsiderable part of marine biodiversity may remain to be diseovered.

Acknowledgements

We are very grateful to A. Kroh, Vienna, for photographing and measuring the speeimens of

O. marmorata, and eounting their spines. We are greatly indebted to G. Paulay and J. Slapeinsky for

aeeess to the FLMNH eolleetion and for eontributing speeimens and images. Our deepest gratitude to

H. Bruggemann who provided all speeimens of O. krohi used in the morphologieal examinations (now

type material) and the eomparative material from the UR eolleetions, and graeiously allowed disseetion

and distribution of types to different institutions. Many thanks also to A. Baldinger, Harvard, for taking

photos of the syntypes of O. insularia, and to C. Ltiter, Berlin, for providing aeeess to the types of

STOHR S. et al.. Taxonomy and phylogeny of the Ophiocoma brevipes group

O. brevipes, O. dentata and O. ternispina. Funding for collection of specimens was provided by the

French Agence Nationale de la Recherche (program BIOTAS, no. ANR-06-BDIV-002) and the National

Science Foundation (NSF DEB-0529724). We thank two anonymous referees for thoughtful comments

that improved the manuscript.

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Manuscript received: 8 July 2013

Manuscript accepted: 26 September 2012

Published on: 9 December 2012

Topic editor: Rudy Jocque

Desk editor: Charlotte Thionois

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT eonsortium: Museum National d’Histoire Naturelle, Paris, Franee; National Botanie Garden

of Belgium, Meise, Belgium; Royal Museum for Central Alfiea, Tervuren, Belgium; Natural History

Museum, Eondon, United Kingdom; Royal Belgian Institute of Natural Seienees, Bmssels, Belgium;

Natural History Museum of Denmark, Copenhagen, Denmark.