Memoirs of Museum Victoria 75:1-5 (2016) Published 2016

1447-2554 (On-line)

http://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

Two new species and a new record of hydroids (hydrozoa: hydroidolina) from Port

Phillip, Australia

Jeanette E. Watson

Honorary Research Associate, Marine Biology Section, Museum Victoria, Melbourne, GPO Box 666E, Victoria, Australia,

email: hydroidw@gmail.com

Abstract Watson, J.E. 2016. Two new species and a new record of hydroids (hydrozoa: hydroidolina) from Port Phillip, Australia.

Memoirs of Museum Victoria 75: 1-5.

A hydroid colony from Port Phillip, southern Australia, yielded two new species, Sertularella eleganta and Bimeria

lutea and a new record of Campanularia laminocarpa Millard, 1966, previously known from South Africa. Four other

known species were epizoic on Sertularella eleganta.

Keywords Southern Australia, Port Phillip, Sertularella eleganta sp. nov., Bimeria lutea sp. nov., Campanularia laminocarpa

Millard, 1966.

Introduction

A collection of hydroids made using scuba from the jetty at the

historic site of South Channel Fort in Port Phillip, southern

Australia, yielded a colony of a new species of Sertularella ( S.

eleganta ), a new species of Bimeria (B. lutea) and a new record

of Campanularia laminocarpa Millard, 1966, previously known

from South Africa. Other species sparsely epizoic on the colony

of S. eleganta were Clytia hemisphaerica (Linnaeus, 1767),

Obelia dichotoma (Linnaeus, 1758), Monotheca flexuosa (Bale,

1894) and Lafoeina amirantensis (Millard and Bouillon, 1973).

The new species and new record are described. Type and

voucher material is lodged in Museum Victoria (NMV F).

Sertularella Gray, 1847

Diagnosis. (Bouillon et al., 2006). Colony erect, branched or

unbranched, monosiphonic or polysiphonic, hydrocaulus and

hydrocladia when present, with two longitudinal rows of

hydrothecae, hydrothecal margin with four cusps, submarginal

teeth present or absent, operculum pyramidal, composed of

four triangular valves, retracted hydranth with abcauline

caecum, gonophores as solitary fixed sporosacs, acrocysts in

some species.

Sertularella eleganta sp. nov.

Figure 1A-F

Material examined. NMV F228240, holotype, colony initially 5%

formalin preserved later transferred to alcohol; fertile colony on rock

in crevice lm deep, coll: J.E.Watson, 22/3/2016. NMV F228241,

microslide malinol mounted, from holotype colony.

Description. Hydrorhiza comprised of narrow stolonal tubes

reptant on concrete surface. Colony without definite main stem,

branching from base, branches monosiphonic except proximally

where some are lightly fascicled from upward-growing stolons

which become primary branches. Branches straight, secondary

branches given off irregularly from primaries below a

hydrotheca at an angle of c. 45°. Branch internodes variable in

length, an indistinct oblique node at junction of adnate and free

hydrothecal adcauline wall, marked by an indentation and

narrowing of perisarc. Proximal internode of secondary branch

cylindrical, long to first hydrotheca.

Hydrothecae alternate, tubular, widely separated along

branches, set at an angle of 40-50° to internodal axis, walls

smooth, narrowing from base to margin. Hydrotheca widest at

junction of adnate and free adcauline wall, adnate adcauline

wall almost parallel to internodal axis, free adcauline wall

slightly concave to straight, ratio of length of adnate to free

adcauline wall 1:2, abcauline wall weakly convex to straight.

Floor of hydrotheca short, transverse to internode with a small

central foramen. Margin delicate with four equidistant cusps

with shallow embayments between and four large thin internal

submarginal cusps of similar shape and size below margin.

Operculum of four very thin flaps. Hydranth too decomposed

for description.

Gonothecae borne abundantly along lower to mid sections

of branches, inserted singly on a short unsegmented pedicel

J.E. Watson

opposite a hydrotheca, facing obliquely upwards. Body of

mature gonotheca elongate oval, variable in length with three

to five broad corrugations, obscure proximally becoming more

prominent distally, surmounted by a long narrow neck above

distalmost deep corrugation, with four equidistant very long,

sharp, often inwardly curved apical spines. Gonophores

female, some extruded from gonotheca as acrocysts.

Perisarc moderately thin throughout. Colour in life pale

yellowish-grey, stolons pale brown.

Table 1. Measurements (pm) of Sertularella eleganta

Branch

intemode length

520-800

width at node

160-184

length to first secondary intemode

700-1000

Hydrotheca

length of abcauline wall

440-480

length of adnate adcauline wall

280-320

length of free adcauline wall

576-650

width at margin

192-208

Gonotheca

length overall

1400-1740

maximum width

680-880

length of neck

296-360

width of neck

168-232

length of apical spines

72-96

Remarks. The colony was growing in a sheltered crevice

between concrete jetty footings in an oceanic strong current-

flow habitat. The delicate flexuous perisarc suggests a deep

water species. Many hydrothecae are infested with one or two

large crustacean eggs.

The nearest congeners of Sertularella eleganta are

Sertularella robusta Coughtrey 1876 and Sertularella

natalensis Millard, 1968. Sertularella. robusta is a very

common southern Australian species occurring in the same

habitat as Sertularella eleganta in Port Phillip. While similar

to S. robusta the hydrothecae of that species are sometimes

faintly rugose, and the gonothecae is more ridged and terminal

spines are shorter. Colony morphology of S. natalensis differs

from S. eleganta in the ratio of fixed:free wall and in striations

on the hydrothecae. Although Sertularella is a genus with

many species, no others have the same morphological,

hydrothecal and gonothecal characters as Sertularella

eleganta.

Etymology. The species name refers to the elegantly branched

colony.

Figure 1A-E. Sertularella eleganta sp. nov. Holotype NMV F228241.

1A fertile branch. IB, part of branch with gonothecae. 1C, branch

internodes. ID, submarginal hydrothecal cusps. IE, gonotheca. Scale

bar: 1A, 20 mm, 1B,C 2 mm, ID, E, 0.5 mm.

Bimeria Wright, 1859

Diagnosis. (Bouillon et al., 2006). Colony stolonal or with

erect branching hydrocauli, stem with firm perisarc enveloping

hydranth, extending as a pseudohydrothecal sheath over

proximal portion of tentacle, hydranth ovoid to vasiform,

hypostome dome-shaped, one or two close whorls of tentacles,

gonophores as fixed sporosacs.

Bimeria lutea sp. nov.

Figure 2A-E

Material examined. NMV F228242 holotype, fertile colony alcohol

preserved, epizoic on Sertularella eleganta, lm deep, coll: J.E.Watson,

22/3/2016. NMV F228243, malinol mounted microslide from holotype

colony.

Description. Colonies fertile, borne abundantly on lower

branches of Sertularella eleganta. Hydrorhiza of tubular

stolons reptant on host colony. Hydrocauli straggling, hydranths

Two new species and a new record of hydroids (hydrozoa: hydroidolina) from Port Phillip, Australia

borne on single pedicels or on sparsely and irregularly branched

stems to 4-5 mm long (rarely 8 mm long). Stems and pedicels

monosiphonic, thick, of same diameter as stolons, deeply

annulated above junction with stolon and above and below each

branch, annulations often fading into corrugations before

becoming smooth. Branching predominantly of first order,

occasionally second order.

Hydrothecae terminal on pedicels of variable length,

hydrotheca vasiform (preserved), a pseudohydrotheca covering

body, hypostome dome-shaped with 10-12 finger-shaped

tentacles arranged in an untidy whorl below hypostome (live

material), the pseudohydrotheca continuing as a thin gelatinous

pellicle over proximal region of tentacles.

Gonophores male, elongate oval, arising singly on a short

annulated pedicel from stem and branches, enclosed in a thick

gelatinous sheath, spadix central, leaf-shaped, opaque.

Cnidome (from live material) clusters of nematocysts of two

categories in transverse bands along tentacles, none discharged:

- microbasic euryteles, loaf-shaped, 4-5 x 8.5-9 pm,

- desmonemes, droplet-shaped, 4x6 pm.

Perisarc very thick on proximal stem region; hydrocaulus,

hydranth and gonophores invested with very fine sediment.

Colour of colony in life: stolons pale brown, hydrocaulus and

tentacles white, hypostome yellow, spadix of gonophore brown.

Table 2. Measurements ( pm) of Bimeria lutea

Stolon, branch width

48-72

Hydranth

length of pedicel

200-2000

length of body

160-180

maximum width

160-184

Gonophore

length of pedicel

64-80

length

360-400

maximum width

112-200

Remarks. Bimeria is a genus of nine species (Bouillon et al.

2006), two of which are known from Australia. Bimeria

australis Blackburn 1937 (redescribed by Watson 1978) is from

the same southern Australian locality as B. lutea, and Bimeria

currumbenensis Pennycuik, 1959 is from tropical southern

Queensland. The morphology of B. lutea fits with neither

Australian species. Colonies of B. australis are not as

abundantly rampant as those of B. lutea-, they are buff-coloured

with a wrinkled hydrocaulus and the stems stand erect from the

substrate. B. currumbenensis described from meagre infertile

material by Pennycuik (1959) is a much larger species and lacks

an annulated hydrocaulus.

Bimeria vestita Wright, 1859 is a known epizoite of

sertulariid hydroids and has been described by Millard (1975),

Calder (1988) and Migotto (1996); the type material was

re-examined by Marques et al. (2000). Differences between

the various descriptions are such that it is likely that more than

one species may be involved. The type as described by

Marques et al. (2000) is much larger and more branched than

B. lutea, the number of tentacles is greater, the nematocysts

are smaller (probably due to shrinkage), the pedicels widen

distally and those of the gonophore are longer. Millard (1975)

mentioned but did not figure a branching male spadix in B.

vestita-, her material may be a different species, possibly more

closely related to B. lutea than to B. vestita.

Etymology. The species is named for the yellow colour of the

hypostome.

Figure 2A-E. Bimeria lutea sp. nov. (holotype colony NMV F228242).

2A, branched stem. 2B, pedicellate hydrotheca (from live material).

2C, male gonophore. 2D, microbasic eurytele. 2E, desmoneme. Scale

bar: 2A, 2 mm. 2B, C, 0.3 mm. 2D, E, 10 pm.

Campanularia Lamarck, 1816

Diagnosis. (Bouillon et al. 2006). Colony stolonal, seldom erect

and branched, hydrorhiza not anastomosing, hydrothecal

pedicel unbranched, hydrotheca campanulate or bell-shaped

with entire or cusped margin, demarcated from pedicel basally

by a variously developed annular perisarcal thickening,

hydrothecal walls with unthickened perisarc, not abruptly

everted distally, true diaphragm absent, subhydrothecal spherule

present, gonophores fixed sporosacs, gonotheca on hydrorhiza.

J.E. Watson

Campanularia laminocarpa Millard, 1966

Figure 3A-E

Campanularia laminocarpa Millard, 1966: 211, fig. 67F-K

Clytia sp. Watson 1975: 158, fig. 1.

Material examined. Microslide NMV F228244, malinol mounted,

from small infertile colony epizoic on Sertularella eleganta in crevice,

lm deep, coll: J.E.Watson, 22/3/2016. Other Material: NMV F228246

microslide, malinol mounted, fertile colony on Synthecium patulum

(Busk, 1852) on reef. North Arm Channel Western Port, 8m, coll: J.E.

Watson 16/12/1996. NMV F228247 microslide, malinol mounted,

fertile colony on Synthecium patulum, reef, 2 km offshore from

McGaurans Beach, Ninety Mile Beach, Bass Strait, 16m, coll: J.E.

Watson 12/8/1983. Microslide (author’s collection), malinol mounted.

Fluted Cape, Tasmania, 15m deep, coll: J.E.Watson, April, 1975.

Description. Colony (NMV F228244) stolonal, hydrorhizal

stolon tubular, reptant on Sertularella eleganta. Hydrocaulus

pedicellate, unbranched, monosiphonic, pedicels variable in

length and width, deeply annulated or spirally ringed

throughout, rarely with smooth patches, pedicel terminating in

a cushion-shaped shoulder supporting a spherule. Hydrotheca

proximally narrow with a moderately long subhydrothecal

chamber with shallow perisarcal distal ring, walls then

widening to become parallel, sometimes expanding, circular in

section. Margin not everted, with 8-10 long cusps separated by

moderately deep and wide embayments, a slight thickening of

perisarc below margin.

Gonothecae male [Western Port (NMV F228246) and

Bass Strait (NMV F228247)], very large, campanulate,

flattened, borne from hydrorhiza on a short unsegmented

pedicel, held obliquely away from host, perisarc smooth

without ornamentation, aperture occupying entire distal

margin, sealed by a thin dome-shaped operculum torn aside at

maturity. Perisarc thin and transparent throughout, gonotheca

fragile and easily collapsed.

Table 3. Measurements ( pm) of Campanularia laminocarpa

Pedicel

length

400-680

width

36-40

Hydrotheca

length overall

368-464

width at margin

128-168

width at diaphragm

44-56

depth of subhydrothecal chamber

36-40

diameter of spherule

40-44

length of cusp

40-48

Gonotheca

length including pedicel

1400-2000

width of margin

800-900

Remarks. I have compared several specimens of

Campanularia epizoic on Synthecium patulum (Busk, 1852)

collected over many years of scuba diving from the southern

Australian localities of Western Port, Bass Strait and

Tasmania with a specimen Campanularia laminocarpa

Millard, 1966 (gift to author from Millard in 1985). Although

the size, shape and dentition of the hydrotheca varies within

Australian localities, morphology and dimensions of the

gonothecae clearly establishes the Australian material as

C. laminocarpa. Minor morphological differences

between the South African and Australian material such as

hydrothecal marginal replication of the South African

species replaced by submarginal thickening in the Australian

material may be due to environmental factors or colony

maturity. The weak perisarcal thickening at the junction

of the subhydrothecal chamber with the body, commented

upon by Millard (1966), is present in some Australian

hydrothecae and can be mistaken for a diaphragm (see

Watson 1975: 158). The hydrothecal margins of the present

specimens of C. laminocarpa are very fragile and easily

collapsed, resulting in changes in apparent shape of the cusps

in mounted specimens.

The small infertile colony on branches of the Sertularella

eleganta host is intergrown with Clytia hemisphaerica.

Figure 3A-E. Campanularia laminocarpa. 3A (NMV F228244),

hydrotheca from Sertularella eleganta , South Channel Fort. 3B,

hydrotheca (NMV F228246) from colony on Synthecium patulum.

North Arm Channel, Western Port. 3C, hydrotheca (NMV F228247)

from colony on Synthecium patulum, off Ninety Mile Beach, Bass

Strait. 3D, E, gonothecae from colony. Ninety Mile Beach, Bass

Strait. Scale bar: 3A-C, 0.3 mm. 3D, E, 1.0 mm.

Two new species and a new record of hydroids (hydrozoa: hydroidolina) from Port Phillip, Australia

References.

Bale, W.M. 1894. Further notes on Australian hydroids with

descriptions of some new species. Proceedings of the Royal

Society of Victoria 2(1): 15-48.

Blackburn, M. 1937. Notes on Australian Hydrozoa, with descriptions

of two new species. Proceedings of the Royal Society of Victoria

50: 170-181.

Bouillon, J., Gravili, C., Pages, F., Gili, J-M. and Boero, F. 2006. An

Introduction to Hydrozoa. Memoires du Museum National

d’Histoire Naturelle 194: 1-591.

Busk, G. 1852. An account of the Polyzoa and sertularian zoophytes

collected on the voyage of the “Rattlesnake” on the coast of

Australia and the Louisiade Archipelago. In MacGillivray. J.

(ed.). Narrative of the voyage of H.M.S. Rattlesnake commanded

by the late Captain O. Stanley during the years 1846-1850 1.

Appendix IV. Boone, London: 343-402.

CalderD.R. 1988. Shallow-water hydroids of Bermuda. The Athecatae.

Life Sciences Contributions of the Royal Ontario Museum 148:

1-107.

Coughtrey, M. 1876. Critical notes on the New Zealand Hydroida.

Transactions and Proceedings of the New Zealand Institute 8:

298-302.

Lamarck, J.B. de 1816. Histoire naturelle des animaux sans vertebres,

vol 2. Verdiere Paris. Pp 1- 568.

Linnaeus, C. 1758. Systerna naturae vol 1, 10 th edition. Holmiae

(Stockholm). L. Salvii, vol. 1: 1-824.

Linnaeus C. 1767. Systema naturae per regna tria naturae: secundum

classes, ordines, genera, species, cum characteribus, differentiis,

synonymis, locis. Editio duodecima. 1. Regnum Animale. 1 & 2

Holmiae, Laurentii Salvii. Holmiae [Stockholm], Laurentii Salvii.

533-1327.

Marques A.C., Mergner H., Hoinghaus R. and Vervoort W. 2000.

Bimeria vestita (Hydrozoa: Anthomedusae: Bougainvilliidae)

senior synonym of Eudendrium vestitum (Hydrozoa:

Anthomedusae: Eudendriidae) Zoologische Mededelingen Leiden

73(22): 321-325.

Migotto A.E. 1996. Benthic shallow-water hydroids (Cnidaria,

Hydrozoa) of the coast of Sao Sebastiao, Brazil, including a

checklist of Brazilian hydroids. Zoologische Verhandlingen

Leiden 306:1-125.

Millard N.A.H. 1966. The Hydrozoa of the south and west coasts of

South Africa. Part III. The Gymnoblastea and small families of

Calyptoblastea. Annals of the South African Museum 48:

427-487.

Millard N.A.H. 1968. South African hydroids from Dr. Th.

Mortensen’s Java-South African expedition 1929-1930.

Videnskabelige Meddelelser fra dansk Naturhistorisk Forening

131: 251-288.

Millard N.A.H. 1975. Monograph on the Hydroids of Southern Africa.

Annals of the South African Museum 68: 1- 513.

Millard N.A.H. and Bouillon, J. 1973. Hydroids from the Seychelles

(Coelenterata). Annales du Musee Royale de TAfrique Centrale,

Serie 8a, Sciences Zoologiques 206:1-106.

Pennycuik PR. 1959. Faunistic records from Queensland. Part V.

Marine and brackish water hydroids. Papers of the Department of

Zoology, University of Queensland 1: 141-210.

Watson J.E. 1975. Hydroids of Bruny Island, southern Tasmania.

Transactions of the Royal Society of South Australia 99(4): 157-

176.

Watson J.E. 1978. New species and new records of Australian athecate

hydroids. Proceedings of the Royal Society of Victoria 90(2):

301-314.

Memoirs of Museum Victoria 75:7-52 (2016) Published 2016

1447-2554 (On-line)

http://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

The sea cucumbers of Camden Sound in northwest Australia, including four new

species (Echinodermata: Holothuroidea)

(http://zoobank.org/urn:lsid:zoobank.org:pub:A7209365-ACCA-4E42-AllF-D211FF09EFD8)

P. MARK O’LoUGHLIN 1 * (http://zoobank.org/urn:lsid:zoobank.org:author:97B95F20-36CE-4A76-9DlB-26A59FBCCE88),

CAROLINE Harding 1 (http://zoobank.org/urn:lsid:zoobank.org:author:FC3B4738-4973-4A74-B6A4-F0E606627674) AND

GUSTAV PAULAY 2 (http://zoobank.org/urn:lsid:zoobank.org:author:A2F155E4-7958-4E63-B36A-CAB23F190A07)

1 Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia

2 Florida Museum of Natural History, University of Florida, Gainesville, FL 32611-7800, USA (paulay@flmnh.ufl.edu)

* To whom correspondence should be addressed. E-mail: pmoloughlin@edmundrice.org

Abstract O’Loughlin P.M., Harding, C. & Paulay, G. 2016. The sea cucumbers of Camden Sound in northwest Australia,

including four new species (Echinodermata: Holothuroidea). Memoirs of Museum Victoria 75: 7-52.

All sea cucumbers collected from Camden Sound by the Kimberley Marine Research Program in 2015 are

reported, with live colour illustrations of the species. Four new species are described, with O’Loughlin as author:

Holothuria ( Metriatyla ) keesingv, Neothyonidium (?) insolitum\ Plesiocolochirus minaeus ; Protankyra torquea.

Colochirus quadrangularis Troschel, the type species of Colochirus Troschel, is reviewed and a sensu stricto diagnosis is

provided for Colochirus. Plesiocolochirus spinosus (Quoy & Gaimard), the type species of Plesiocolochirus Cherbonnier,

is reviewed and a sensu stricto diagnosis is provided for Plesiocolochirus. Colochirus robustus Ostergren is confirmed for

NW Australia, but not for Camden Sound. Pseudocolochirus axiologus (H. L. Clark) is raised out of synonymy with

Pseudocolochirus violaceus (Theel). Thyone papuensis Theel is reported from Camden Sound and the species is reviewed

and illustrated. We report Thyone pedata Semper from Joseph Bonaparte Gulf in northern Australia, but not for Camden

Sound. The WA Naturalists Club visited “Camden Harbour” in 1990 and Marsh reported on the marine invertebrates. Two

sea cucumber species from this report are included here. A phylogenetic tree is provided with sequences for species of

Colochirus and Plesiocolochirus. A table is provided with a list of all sea cucumbers collected from Camden Sound.

Tissue samples for genetic analysis were taken from all specimens, and tissue data are listed in two tables. Two Pilumnidae

crabs were found in the coelom of the new species Plesiocolochirus minaeus.

Keywords Kimberley; Camden Sound; Colochirus ; Metriatyla ; Neothyonidium; Pilumnidae; Plesiocolochirus ; Protankyra ;

Pseudocolochirus; Thyone.

Introduction

Camden Sound, in the Kimberley Region of northwest Western

Australia, is southwest of Augustus Island (-15.40 124.63) and

west of Kuri Bay and Brecknock Harbour (“Camden

Harbour”). In 2012 the Western Australia State Government

created the Camden Sound Marine Park. Subsequently the

Western Australian Marine Science Institution informs the

management and monitoring of the Region through the

Kimberley Marine Research Program. A ship-based

expedition to Camden Sound was conducted in March 2015

under the auspices of WAMSI’s Kimberley Benthic

Biodiversity Project using AIMS’s RV Solander and CSIRO’s

RV Linnaeus. All holothuroid echinoderm (sea cucumber)

specimens were sent to Museum Victoria for determination,

and the collection is the subject of this report. Colour

photographs of live sea cucumber specimens were taken

during the KMRP expedition, principally by John Keesing

(CSIRO), and photographs of all the species are published in

this work. The Camden Sound sea cucumber collections are

lodged in the Western Australian Museum.

The WA Naturalists Club visited “Camden Harbour” in

1990. Marsh (2011) reported on the marine invertebrates.

Holothuroid species from Adele Island and Montgomery Reef

were reported but both locations are remote from Camden

Sound. The two species collected from Slate Island at the

southern edge of Camden Sound are included in this work:

Holothuria ( Halodeima ) atra Jaeger, 1833; Holothuria

(. Mertensiothuria ) leucospilota (Brandt, 1835).

We note that the ICZN (Opinion 417,42 pp., 1956) rejected

for nomenclatorial purposes the publication by Oken 1815, and

as a consequence the genera Psolus Oken, 1815 and Thyone

Oken, 1815 became invalid. The Commission has now ruled in

P.M. O’Loughlin, C. Harding & G. Paulay

Table 1. Sea cucumber species collected from Camden Sound.

Order

Family

Subfamily

Taxon

Aspidochirotida

Holothuriidae

l Holothuria (Halodeima) atra Jaeger, 1833 (Slate Island; WAM Z58692)

1 Holothuria (Mertensiothuria) leucospilota (Brandt, 1835) (Slate Island;

WAMZ58735)

Holothuria ( Metriatyla ) keesingi O’Loughlin sp. nov.

Holothuria ( Thymiosycia ) gracilis Semper, 1868

Stichopodidae

Stichopus unresolved species complex including Stichopus herrmanni

Semper, 1868

Dendrochirotida

Cladolabidae

Globosita elnazae O’Loughlin, 2014 (in O’Loughlin, Mackenzie &

VandenSpiegel 2014)

Cucumariidae

Colochirinae

Cercodemas anceps Selenka, 1867

Colochirus quadrangular is Troschel, 1846

Leptopentacta grisea H. L. Clark, 1938

Plesiocolochirus sp. 1, unresolved species complex including P. australis

(Ludwig, 1875)

Plesiocolochirus minaeus O’Loughlin sp. nov.

Pseudocolochirus axiologus (H. L. Clark, 1914)

Phyllophoridae

Phyllophorus (Urodemella) holothurioides Ludwig, 1875

Phyllophorella spiculata (Chang, 1935)

Sclerodactylidae

Havelockia versicolor (Semper, 1867)

Thyonidae

Semperiellinae

Massinium bonapartum O’Loughlin, 2014 (in O’Loughlin, Mackenzie &

VandenSpiegel, 2014)

Neothyonidiumip .) insolitum O’Loughlin sp. nov.

Thyoninae

Hemithyone semperi (Bell, 1884)

Stolus canescens (Semper, 1867)

Thyone papuensis Theel, 1886

Thyonidiidae

Actinocucumis longipedes H. L. Clark, 1938

Actinocucumis solanderi O’Loughlin, 2014 (in O’Loughlin, Mackenzie &

VandenSpiegel, 2014)

Actinocucumis typica Ludwig, 1875

Mensamaria intercedes (Lampert, 1885)

Molpadida

Molpadiidae

Molpadia scabrum (Sluiter, 1901)

Synaptida

Synaptidae

Rynkatorpinae

Protankyra insolens (Theel, 1886)

Protankyra torquea O’Loughlin sp. nov.

Protankyra verrilli (Theel, 1886)

Synaptinae

Synaptula lamperti Heding, 1928

Synaptula recta (Semper, 1867)

WAM specimens collected at Slate Island by WANC in July 1990, and reported by Marsh (2011).

favour of their availability (Opinion 2367) in response to an

application to the ICZN by Paulay & O’Loughlin (Case 3598)

for both Psolus Oken, 1815 and Thyone Oken, 1815 to be made

available.

Abbreviations

AIMS Australian Institute of Marine Science

CSIRO Commonwealth Scientific and Industrial Research

Organization

GA Geoscience Australia

KMRP

LKCNHM

MOL AF

MRAC

NMV

NUS

PMCP

Kimberley Marine Research Project

Lee Kong Chian Natural History Museum

Prefix for code number of tissues provided to the

University of Florida for sequencing

Royal Museum for Central Africa, Tervuren

Museum Victoria, with specimen registration

prefix F

National University of Singapore

University of the Philippines

Pilbara Marine Conservation Program

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

TMSI Tropical Marine Science Institute (Singapore)

UF University of Florida

USNM United States National Museum (Smithsonian

Institution)

WAM Western Australian Museum, with specimen

registration prefix Z

WAMSI Western Australian Marine Science Institution

WANC The Western Australian Naturalists Club

ZRC Zoological Reference Collection of LKCNHM

ZRC.ECH LKCNHM catalogue number prefix (echinoderms)

Methods.

All specimens were preserved in 100% ethanol on the vessel

by WAM staff, and databasing and weighing of specimens was

done by CSIRO. The colour photos of live specimens published

here were taken at the time of collection, principally by John

Keesing (CSIRO) using a Nikon D300 digital SLR camera.

Some specimens were photographed without a scale. We have

estimated that there is about 25% shrinkage of soft-bodied

specimens when preserved in 100% ethanol, and no shrinkage

in hard-bodied specimens. We have thus been able to provide

an estimated live colour size in the captions when there is no

scale bar for the live photos of the now preserved specimens.

Most of the macro images of preserved specimens were taken

by Caroline Harding, with Mark O’Loughlin, using a Canon

5D mark ii camera mounted on a camlift Visionary Digital

auto stepper. A Zerene Image Stacker, Adobe Lightroom and

Photoshop were used for image processing and editing. Macro

images of the preserved holotype of Neothyonidium(l )

insolitum were taken by Melanie Mackenzie (NMV) with a

Leica DC500 high resolution digital camera system with Auto

Montage software. The photos of ossicles were taken by

Caroline Harding, with Mark O’Loughlin, using a LEICA

DM5000 B microscope, Leica application software, and

Helicon Focus montage software.

Tissues were sent to Gustav Paulay (UF) for sequencing,

and specimen source locations, tissue codes, catalogue

numbers and GenBank Accession numbers are recorded in

Appendices 1 and 2. A 655 bp portion of the mitochondrial

gene cytochrome oxidase subunit 1 (COI) was sequenced from

selected specimens using the echinoderm barcoding primers

COIceF (5’-ACTGCCCACGCCCTAGTAATGATATTTTTT

ATGGTNATGCC-3’) and COIceR (5’ TCGTGTGTCTACGT

CCATTCCTACTGTRAACATRTG-3’) (Hoareau & Boissin

2010), as described in Michonneau & Paulay 2014. We note

that these echinoderm specific primers amplify positions

242 to 898 in COI compared with positions 74 to 733 amplified

by Folmer primers. COI sequences were aligned by eye a

nd analyzed using Maximum Likelihood with 100

bootstrap replicates, implemented in MEGA 6.06 (Tamura et

al. 2013). Sequences have been submitted to GenBank (See

Appendix 2).

Terminology.

For small concave plates, with two large central and two smaller

distal perforations, and sometimes with additional small outer

perforations, we use the term bowl, not cup or basket.

Order Aspidochirotida Grube, 1840

Holothuriidae Burmeister, 1837

Holothuria (.Metriatyla ) Rowe, 1969

Holothuria ( Metriatyla) keesingi O’Loughlin sp. nov.

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act:

ED316CAC-695D-4EB7-97DD-8999F2CD33CF

Table 1; appendix 1; figures 2a, b, 3, 4

Material examined. Holotype. Northwest Western Australia,

Kimberley Region, Camden Sound, WAMSI 1.1.1, RV Solander, sled,

site no SOL_47, WAM station no 42, barcode 10002938, from

-15.612805 124.073033 36 m to -15.612437 124.072883 35 m, 26 Mar

2015, WAM Z89006.

Paratypes. Camden Sound, WAMSI 1.1.1, RV Solander , sled, site

no SOL_107, WAM station no 1, barcode 10000043, from -15.514826

124.183111 46 m to -15.514503 124.183774 45 m, 14 Mar 2015, WAM

Z89000 (1 specimen); RV Solander, sled, site no SOL_8, WAM

station no 7, barcode 10000403, from -15.313929 124.112518 47 m to

-15.313336 124.111992 48 m, 16 Mar 2015, WAM Z89001 (3); RV

Solander, sled, site no LIN_36, WAM station no 17, barcode 10001168,

from -15.220444 124.320894 50 m to -15.220159 124.320648 50 m,

18 Mar 2015, WAM Z89002 (2); RV Solander, sled, site no SOL_32,

WAM station no 19, barcode 10001320, from -15.253592 124.203038

45 m to -15.253318 124.202302 45 m, 19 Mar 2015, WAM Z89003 (1);

RV Solander, sled, site no SOL_24, WAM station no 23, barcode

10001821, from -15.40642783 124.1259284 42 m to -15.40693704

124.1253687 42 m, 20 Mar 2015, WAM Z89004 (2); RV Solander,

sled, site no SOL_87, WAM station no 24, barcode 10001954, from

-15.448727 124.153629 36 m to -15.44933 124.154105 36 m, 20 Mar

2015, WAM Z89005 (2); RV Solander, sled, site no SOL_47, WAM

station no 42, barcode 10002963, from -15.612805 124.073033 36 mto

-15.612437 124.072883 35 m, 26 Mar 2015, WAMZ89007 (1).

Description (preserved in 95% ethanol). Up to 60 mm long,

18 mm wide, 12 mm high; body surface finely nodulose; body

arched dorsally, with rounded ventro-lateral margins, low convex

ventrally; strongly tapered anteriorly and posteriorly; dorsal and

lateral papillae irregularly distributed, conical, with tapered to

pointed ends, of variable sizes, up to 3 mm long; about eight

papillae across body transversely, longest on dorso-lateral radii,

about 40 ventro-lateral papillae in close irregular series on each

margin; tube feet digitiform, up to 2 mm long, scattered on

ventrum but in recognizable irregular longitudinal series, paired

irregular series latero-ventrally, paired irregular series on each

side of bare mid-ventrum; mouth antero-ventral, with 20 tentacles,

mouth surrounded by a ring of about 16 conical papillae, up to

1 mm long; calcareous ring solid, widths of radial and inter-radial

plates sub-equal, inter-radials half the height of the radials, with

undulating posterior edge, lacking posterior prolongations.

Body wall ossicles large tables and buttons, buttons more

abundant than tables; table discs of variable size, shallow

concave, irregularly round to rounded square to oval, margin

smooth, discs 48-240 pm across, disc perforations from 8 to

more than 50, perforations very small marginally; table spires

of variable height, up to 176 pm long, 4 pillars, up to 8 cross

bridges, spire rounded distally with cluster of small spinelets,

sometimes spinelets extend along distal sides of spire; buttons

P.M. O’Loughlin, C. Harding & G. Paulay

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Figure 1. Maximum likelihood tree of Colochirus-Plesiocolochirus COI data, with mid-point rooting. Bootstrap support (100 replicates)

indicated by circles (100%) and rectangles (>95%).

C quadrangularis

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 2. a, b, photos of live holotype specimen of Holothuria ( Metriatyla ) keesingi O’Loughlin sp. nov. (WAM Z89006): a, dorso view; b,

latero-ventral view with ventrum and tube feet along upper side, dorsal papilla underneath, c, photo of dorsal view of live specimen of Holothuria

(Thymiosycia) gracilis Semper, 1868 (WAM Z89008; estimated 125 mm long live).

P.M. O’Loughlin, C. Harding & G. Paulay

Figure 3. Preserved holotype of Holothuria ( Metriatyla) keesingi O’Loughlin sp. nov. (WAM Z89006): a, dorsal view; b, ventral view; c,

tentacles with surrounding ring of papillae; d, calcareous ring with radial plate right, inter-radial plate left.

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

mrM

Figure 4. Ossicles from holotype (WAM Z89006; rods and small button mid-top and bottom) and paratype (WAM Z89005; tables, buttons)

specimens of Holothuria ( Metriatyla ) keesingi O’Loughlin sp. nov. (in specimens table discs up to 240 m across; table spires up to 176 pim long;

buttons up to 200 /*m long; rods up to 336 }im long).

P.M. O’Loughlin, C. Harding & G. Paulay

predominantly elongate with up to about 7 irregularly paired

perforations, some buttons with 1-3 pairs of perforations, button

sizes 50-200 pm long, buttons smooth and variably knobbed.

Dorsal papillae with tables, buttons and thick perforated rods;

rods thick, mid-rod widened with perforations, distal ends

widened with small perforations, rods up to 336 p m long.

Ventral tube feet with endplates, thick endplate support rods,

tables and buttons; endplates slightly convex, central perforations

slightly larger, margin bluntly digitiform, endplate diameters

about 300 ja m; thick rods curved, widened and perforated mid¬

rod and distally, up to 264 jAm long. Tentacles with minutely

spinous, non-perforate, curved rods, rods up to 320 jAm long.

Colour (preserved). Body pale mottled brown to off-white,

dorsal and lateral irregular fine brown-black flecks and spots,

about 4 dorso-lateral pairs of large irregular brown-black

patches that are partly merged at the anterior end; tentacles and

distal papillae and tube feet yellow. Live colour similar but the

body base colour yellow.

Distribution. NW Western Australia, Kimberley Region,

Camden Sound, mud, 35-50 m.

Etymology. Named for John Keesing of the CSIRO Oceans and

Atmosphere, and The Western Australian Marine Science

Institution, with appreciation of his live colour photography of

Camden Sound sea cucumbers used in this work, and his

gracious collaboration with data.

Remarks. The morphology of Holothuria (Metriatyla) keesingi

O’Loughlin sp. nov. satisfies the diagnosis of the Holothuria

sub-genus Metriatyla Rowe, 1969:

1. 20 tentacles;

2. collar of papillae around the base of the tentacles;

3. large conical, irregularly arranged papillae dorsally, a

lateral flange sometimes evident;

4. tube feet irregularly arranged on the ventrum;

5. body arched dorsally, flattened ventrally;

6. size small to large;

7. body wall thin to thick;

8. calcareous ring well developed;

9. table ossicles with smooth disc and spire of moderate height

to high, terminating in a few to many small spines;

10. buttons simple, with moderate-sized irregularly arranged

knobs and 3-10 pairs of relatively large holes.

We qualify the diagnostic characters of Rowe (1969) to include

the possibility of species with a large body and thick body wall

(as in Holothuria (Metriatyla) scabra Jaeger, 1833).

Rowe (in Rowe & Gates 1995) synonymised Holothuria

bowensis Ludwig, 1875 (type from Bowen, NE Australia) and

Holothuria subverta H. L. Clark, 1921 (type from Torres

Strait, NE Australia) with Holothuria (Metriatyla) martensii

Semper, 1868 (type from Amboina, Indonesia). Of the species

referred to sub-genus Metriatyla, the new species resembles

Holothuria (Metriatyla) martensii. However, the new species

Holothuria (Metriatyla) keesingi O’Loughlin is significantly

different in the following ways:

1. smaller species with numerous preserved specimens up to

only 60 mm long;

2. preserved colour with yellow papilla and tube foot and

tentacle ends, and dorsal body with four irregular longitudinal

paired dark brown patches;

3. larger tables, with discs up to more than 200 pm across, disc

perforations up to more than 40, spires up to more than

150 pm high;

4. larger smooth and knobbed buttons, predominantly about

7 irregular pairs of perforations, up to 200 pm long.

Theel (1886) referred two specimens to Holothuria

(Metriatyla) martensii, one from Indonesia and one from the

Philippines. Sizes were 150 mm and 85 mm long. Buttons

were up to 140 pm long. Table disc perforations were fewer

than 30. Colours are described with no reference to brown

dorsal patches. These morphological characters of Holothuria

(Metriatyla) martensii (sensu Theel 1886) are significantly

different to those of Holothuria (Metriatyla) keesingi

O’Loughlin sp. nov.

The buttons in H. (Metriatyla) horrida Massin, 1987 are

similar to those in Holothuria (Metriatyla) keesingi

O’Loughlin. And the high table spires of H. (Metriatyla)

horrida are also similar but significantly shorter (up to 120 pm

long). The table discs in H. (Metriatyla) horrida are

significantly smaller (up to 140 pm across), the table disc

perforations significantly fewer (up to 16), and the colour

reported as grey.

Frank Rowe (pers. comm) suggested that the size and

form of the ossicles in the relatively small specimens appear

somewhat paedomorphic. But the mature gonads that are

present confirm that the specimens are adult.

Holothuria (Thymiosycia) gracilis Semper, 1868

Table 1; appendix 1; figure 2c

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, Ngalanguru Island, 25 Mar 2015, WAM Z89008 (1).

Remarks. This species identity was established by Fran§ois

Michonneau (U F,pers. comm).

Stichopodidae Haeckel, 1896

Stichopus sp.

Table 1; appendix 1; figure 5

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, Montgomery Island reef flat, barcode 10003037, 24

Mar 2015, WAM Z89009 (1); barcode 10003038, 24 Mar 2015, WAM

Z89010 (1).

Remarks. The taxonomy of Stichopus is challenging today.

Based on the living appearance of this animal it could represent

Stichopus horrens Selenka, 1867, or a specimen in the

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 5. Photos of live specimen belonging to a Stichopus unresolved species complex (WAM Z89009; estimated 110 mm long live): a, dorsal

view; b, ventral view; c, in situ view.

P.M. O’Loughlin, C. Harding & G. Paulay

S. monotuberculatus (Quoy & Gaimard, 1834) or S. herrmanni

Semper, 1868 species complexes. Specimens from NW

Australia identified by Mark O’Loughlin previously as

Stichopus herrmanni are now revised to “ Stichopus unresolved

species complex”.

Order Dendrochirotida Grube, 1840

Family Cladolabidae Heding & Panning, 1954 sensu Smirnov

2012

Remarks. See O’Loughlin, Mackenzie & VandenSpiegel 2014.

Globosita Cherbonnier, 1958

Remarks. See O’Loughlin, Mackenzie & VandenSpiegel 2014.

Globosita elnazae O’Loughlin, 2014 (in O’Loughlin,

Mackenzie & VandenSpiegel 2014)

Table 1; appendix 1; figures 6a, b

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no SOL_116a,

WAM station no 18, barcode 10001261, from -15.261423 124.275183

41 m to -15.261716 124.275827 40 m, 18 Mar 2015, WAM Z89011 (1);

same data, barcode 10001262, WAM Z89012 (1).

Family Cucumariidae Ludwig, 1894

Diagnosis (after Smirnov 2012). Ten dendritic tentacles;

calcareous ring lacking segmented posterior prolongations;

tube feet most commonly restricted to the radii, or may also be

scattered in the dorsal and lateral inter-radii; ossicles in body

wall perforated plates, sometimes rods, sometimes bowls,

never tables.

Subfamily Colochirinae Panning, 1949

Diagnosis. Cucumariidae with plate and bowl ossicles.

Cercodemas Selenka, 1867

Cercodemas Selenka, 1867: 343.—Rowe (in Rowe & Gates),

1995: 271.

Remarks. Rowe (in Rowe & Gates 1995) raised genus

Cercodemas out of synonymy.

Cercodemas anceps Selenka, 1867

Table 1; appendix 1; figures 6c, d

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no SOL_43,

WAM station no 26, barcode 10002087, from -15.488461 124.201824

46 m to -15.488309 124.201113 46 m, 21 Mar 2015, WAM Z89013 (1).

Remarks. We note that in relation to Colochirus Troschel, 1846

(below) and Plesiocolochirus Cherbonnier, 1946 (below), the

monotypic Cercodemas Selenka, 1867 has body wall ossicles

comprising deep bowls, deep bowls bridged over rim to create

hollow ellipsoids, and multi-layered scales, but lacks shallow

bowls and knobbed buttons.

Colochirus Troschel, 1846

Figure 1; appendices 1, 2

Colochirus Troschel, 1846: 64.—Semper, 1867: 56.—Ekman,

1918: 5-6.—Panning, 1949: 439.—Panning, 1971: 42-43, fig. 5.—Liao

& Clark, 1995: 474.—Rowe (in Rowe & Gates), 1995: 272.

Type species. Colochirus quadrangularis Troschel, 1846

(monotypy)

Other currently assigned, accepted species (with type locality

added). Colochirus crassus Ekman, 1918 (NW Australia; junior

synonym Colochirus quadrangularis var. australoides Panning,

1949 by Rowe in Rowe & Gates 1995); C. cylindricus Semper,

1867 (Philippines); C. pusillus Heifer, 1912 (Gulf of Suez); C.

robustus Ostergren, 1898 (S Korea; junior synonym Colochirus

squamatus Sluiter, 1901 by Rowe in Rowe & Gates 1995).

Diagnosis (sensu stricto, based on type species only, described

below). Body quadrangular in mid-body section, slightly

tapered towards oral and anal ends; body and papillae firm,

densely packed with ossicles; preserved length up to 98 mm,

dorsal and ventro-lateral radii slightly raised, each with

prominent, conical papillae in irregular zig-zag rows; five oral

valves, each with a terminal papilla and sometimes 1 or 2

additional papillae; five anal scales, some small peri-anal

papillae, 5 longer proximal anal radial papillae. Ten dendritic

tentacles, 8 large, 2 ventral small. Calcareous ring plates not

forked posteriorly, lacking posterior prolongations. Dorsal and

lateral inter-radii lacking tube feet; tube feet in discrete bands

in ventro-lateral and mid-ventral radii, each band about 4 podia

wide, ventral inter-radii usually lacking tube feet, inter-radii

similar in width to the radial bands of tube feet.

Ossicles of body wall of six recognizable but inter-grading

forms:

1. small, shallow bowls, margin smooth to finely knobbed or

finely spinous, variably bridged across margins or not;

2. abundant rounded-rectangular to oval to irregular shallow

bowls, one short or long margin prominently spinous, bowls

variably partly bridged or not;

3. some rounded-rectangular to oval larger shallow bowls,

sometimes partly bridged; 4. larger shallow bowls bridged on

one side to create smooth, hollow, irregular ellipsoids;

5. shallow bowls, frequently thick-walled, bridged to create

irregular ellipsoids with inner bridging, not hollow;

6. enlarged inner-bridged ellipsoids inter-grade with multi¬

layered scales.

Tentacle ossicles rod-plates, rods, rosettes.

Remarks. Rowe (in Rowe & Gates 1995) restricted Colochirus to

three assigned species Colochirus crassus, C. quadrangularis

and C. robustus, leaving the status of C. cylindricus, C. pusillus

and C. viridis Semper, 1867 (considered to be nomen dubium ) as

uncertain, as they were not assigned to other genera.

With the support of COl sequence data (Figure 1), we

confirm Ekman’s report (1918) of the occurrence of Colochirus

robustus in NW Australia.

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 6. a, b, photos of live specimens of Globosita elnazae O’Loughlin, 2014 (in O’Loughlin, Mackenzie & VandenSpiegel 2014): a, lateral

view (WAM Z89011; estimated 25 mm long live); b, posterior dorsal view (WAM Z89012; estimated 90 mm long live), c, d, photos of live

specimen of Cercodemas anceps Selenka, 1867 (WAM Z89013): c, dorsal view; d, ventral view.

P.M. O’Loughlin, C. Harding & G. Paulay

A phylogenetic analysis based on COl sequence data

shows that species currently assigned to Colochirus and

Plesiocolochirus fall in two well-supported clades (Figure 1).

The Colochirus cluster includes Colochirus quadrangularis

(from Singapore, near the type locality, and across tropical

Australia), Colochirus robustus (from NW Australia and the

Philippines), and Colochirus species 1 (from N Australia,

Japan, the Philippines, and Madagascar). This unassigned

species is morphologically close to specimens that have

frequently been identified as Plesiocoloshirus australis

(Ludwig, 1875). After examining type material, Rowe (in

Rowe & Gates 1995) synonymised Colochirus minutus

Ludwig, 1875 with Plesiocolochirus australis (Ludwig, 1875).

We hope to examine the type specimens of both of these

species and then be in a position to assign the unidentified

Colochirus species in a subsequent work.

Panning (1971) observed that some shallow bowls (up to

107 pm long) were bridged on both sides to create irregular

ellipsoids with an inner shallow bowl layer, and thus not

hollow. We have not been able to support this observation by

Panning (1971) as all solid ellipsoids appear to be built up on

the concave side only of the shallow bowls with the outer

surface supported by an inner network. Shallow bowls bridged

on one side only but without an inner bridging network create

irregular hollow ellipsoids. We note that Cercodemas anceps

Selenka, 1867 has deep bowls that may be bridged to create

such more regular hollow ellipsoids.

Colochirus Troschel, 1846 (type species Colochirus

quadrangularis Troschel, 1846, above) is distinguished sensu

stricto from Plesiocolochirus Cherbonnier, 1946 (type species

Plesiocolochirus spinosus (Quoy & Gaimard, 1834), below), by:

1. absence of large imbricating external scales on distal anal

cone, on proximal oral valves, and on lateral papillae;

2. absence, usually, of inter-radial tube feet;

3. absence of knobbed single-layered button ossicles in the

body wall;

4. presence of tentacle rosettes.

Colochirus quadrangularis Troschel, 1846

Table 1; appendices 1, 2; figures 1, 7, 8, 9

Colochirus quadrangularis Troschel, 1846: 64-66 ( non

Holothuria quadrangularis Lesson, 1830: 90-91, pi. 31 fig. 1).—

Theel, 1886: 81-82, 120-121, pi. 6 fig. 7, pi. 14 figs 7, 8.-Erwe, 1913:

353-355, fig. 2a-g.—Ekman, 1918: 21-26, pi. 1 figs 7-10, pi. 3 figs

13-15.—Panning, 1949: 446-447, figs 46, 47,-Liao & Clark, 1995:

474-475, fig. 286.—Rowe (in Rowe & Gates), 1995: 272-273.

Colochirus coeruleus Semper, 1867: 59, pi. 11 fig. 1, pi. 13 fig. 18

(synonymy by H. L. Clark 1946).

Colochirus jagorii Semper, 1867: 60.—Panning, 1971: 42 (type

locality Singapore; synonymy by Rowe (in Rowe & Gates) 1995).

Colochirus tristis Ludwig, 1875: 87-88 (type locality Zanzibar;

synonym of Colochirus jagorii by Panning 1971).

Pentacta quadrangularis.— H. L. Clark, 1946: 391.—Cannon &

Silver, 1986: 30.

Pentacta coerulea.—H. L. Clark, 1932: 227—H. L. Clark, 1938:

449-450, pi. 16 fig. 4.

Pentacta jagorii— H. L. Clark, 1932: 228-229.—H. L. Clark,

1946: 391-392.

Pentacta coerulea var, rubra H. L. Clark, 1938: 451, pi. 16 fig. 5

(single specimen from Broome, NW Australia)

Type locality. Coast of Malacca (southern region of the Malay

Peninsula, near Singapore).

Material examined. Singapore, Johor Strait, dredge, channel between

Beting Bronok and Chek Jawa channel, 1.41 103.98 1.5-2.4 m, mud,

coll. Tan Koh Siang et al. , 29 Jun 2011, NMV F210388 (1) (former

registration ZRC.ECH.0208; donated to Museum Victoria); west end

of Jurong island (composite new island), 1.22 103.67 23.1-24.4 m,

dredge, rock, sand and mud, coll. Lim Swee Cheng et al., 19 Dec 2013,

NMV F210389 (1) (former registration SEA-3046; donated to

Museum Victoria).

Northwest Western Australia, Kimberley Region, Camden Sound,

WAMSI 1.1.1, RV Solander, sled, site no SOL_107, WAM station no

1, barcode 10000001, from -15.514826 124.183111 46 m to -15.514503

124.183774 45 m, 14 Mar 2015, WAM Z89014 (1); RV Solander, sled,

site no SOL_60, WAM station no 9, barcode 10000582, from

-15.311436 124.162869 42 m to -15.311705 124.162473 42 m, 16 Mar

2015, WAM Z89015 (1); RV Solander, sled, site no SOL_60, WAM

station no 9, barcode 10000647, from -15.311436 124.162869 42 m to

-15.311705 124.162473 42 m, 16 Mar 2015, WAM Z89016 (1); RV

Solander, sled, site no LIN_36, WAM station no 17, barcode 10001114,

from -15.220444 124.320894 50 m to -15.220159 124.320648 50 m, 18

Mar 2015, WAM Z89017 (5); RV Solander, sled, site no SOL_32,

WAM station no 19, barcode 10001331, from -15.253592 124.203038

45 m to -15.253318 124.202302 45 m, 19 Mar 2015, WAM Z89018 (6);

RV Solander, sled, site no SOL_56, WAM station no 20, barcode

10001420, from-15.376537 124.192773 35 m to-15.376196 124.192071

35 m, 19 Mar 2015, WAM Z89019 (1); RV Solander, sled, site no

SOL_84, WAM station no 21, barcode 10001560, from -15.414697

124.059193 36 m to -15.415001 124.059918 36 m, 20 Mar 2015, WAM

Z89020 (1); RV Solander, sled, site no SOL_24, WAM station no 23,

barcode 10001767, from -15.406428 124.125928 42 m to -15.406937

124.125369 42 m, 20 Mar 2015, WAM Z89021 (1); RV Solander, sled,

site no SOL_160, WAM station no 25, barcode 10001971, from

-15.428534 124.273164 17 mto -15.428908 124.273525 17 m, 21 Mar

2015, WAM Z89022 (2); RV Solander, sled, site no SOL_109a, WAM

station no 33, barcode 10002500, from -15.711677 124.2303 25 m to

-15.711054 124.229793 25 m, 23 Mar 2015, WAM Z89023 (1); RV

Solander, sled, site no SOL_73, WAM station no 38, barcode

10002692, from -15.945442 124.366373 29 m to -15.945268

124.367171 28 m, 25 Mar 2015, WAM Z89024 (1); RV Solander, sled,

site no SOL_69, WAM station no 41, barcode 10002871, from

-15.747648 124.146502 43 m to -15.747285 124.14634 43 m, 26 Mar

2015, WAM Z89025 (1).

Joseph Bonaparte Gulf, cruise SOL 4934, station 33, sample

29024, -11.65 129.83 24 m, NMV F201791 (11) (tissue code MOL AF

1516); NMV F173258; NMV F173259; NMV F173260; NMV

F173261; NMV F201781; NMV F201782; NMV F201788; NMV

F201789; NMV F201790; NMV F201792; NMV F203004

Other Northern Australia, NMV F95253; NMV F95254; NMV

F95255; NMV F95256; NMV FI 12191; NMV FI 12192; NMV

FI 13573; NMV FI 13574; NMV FI 13575; NMV FI 13576;

NMVF113577; NMVF113578; NMV FI 13579; NMV FI 13580; NMV

FI 13581; NMV F149742.

Great Australian Bight, NMV FI 13582; NMV F199464.

Description. Body quadrangular in section, slightly tapered

towards oral and anal ends, anal end slightly upturned,

preserved body (excluding tentacles) up to 98 mm long, body

surface with tessellated appearance or smooth; body and

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 7. Photos of live specimens of Colochirus quadrangularis Troschel, 1846 from Singapore waters, provided by Helen Pei San Wong and

Joo Yong Ong (TMSI of NUS; specimens estimated to be up to 60 mm long): a, dorsal view showing anal scales and absence of warts; b, ventral

view; c, dorsal view showing warts; d, lateral view.

P.M. O’Loughlin, C. Harding & G. Paulay

Figure 8. Photos of live specimens of Colochirus quadrangularis Troschel, 1846 from northern Australia: a, dorsal view (WAM Z89021, from

Camden Sound); b, ventral view (WAM Z89015, from Camden Sound); c, colour morphs from Joseph Bonaparte Gulf (NMV F201791).

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 9. Ossicles from specimens of Colochirus quadrangularis Troschel, 1846 from Singapore: a, dorsal and peri-anal body wall bowls up to

56 pim long (three with single bridge) (top row), bowls with one spinous margin up to 80 pim long (three with single bridge) (middle row), hollow

ellipsoids up to 88 pim long, and ellipsoids with inner bridges up to 88 pim long (bottom row) (from Singapore specimen NMV F210388); b,

tentacle rods and rod-plates up to 440 pim long, rosettes up to 96 pim long (from NMV F210388); c, ventral tube foot endplate up to 400 pim

diameter (right), support rod-plates up to 272 pim long, small bowls, spinous-edge bowl with bridging, and ellipsoid with internal bridging

(bottom left) (from NMV F210389).

P.M. O’Loughlin, C. Harding & G. Paulay

papillae firm, densely packed with ossicles; dorsal and ventro¬

lateral radii slightly raised, each with about 12 conical papillae

in irregular zig-zag rows, papilla lengths variable up to about

10 mm long, papillae variably straight to curved, surmounted

by tube feet ventro-laterally and sometimes dorsally, anterior

and posterior ventral radii all with papillae; dorsal and lateral

inter-radii slightly depressed, variably with or lacking scattered

short conical or wart-like protuberances, lacking tube feet;

5 anterior oral valves, each with a terminal papilla and

sometimes 1 or 2 additional papillae; 5 inner anal scales, some

small peri-anal papillae, 5 longer proximal anal radial papillae;

10 dendritic tentacles, ventral 2 smaller; calcareous ring plates

not forked posteriorly and lacking posterior prolongations; tube

feet in discrete bands on ventral radii, each band about 4 wide,

discrete inter-radii usually lacking tube feet, ventral inter-radii

similar in width to the radial bands of tube feet.

Intergrading ossicle forms of dorsal mid-body wall (from

NMV F210388):

1. on body wall surface, irregular oval to rounded-rectangular

shallow bowls, long margins sometimes indented, four large

central perforations, usually four small corner perforations,

sometimes additional smaller marginal perforations, rim variably

smooth or with fine knobs or fine blunt spines, bowls with or

lacking bridges across rim, bowls up to rarely 56 pm long;

2. outer body wall, abundant rounded-rectangular to oval to

irregular shallow bowls, four large central perforations, smaller

peripheral perforations, one short or long margin prominently

spinous, bowls variably partly bridged or not, up to 80 pm long;

3. inner body wall, some rounded-rectangular to oval shallow

bowls, 4 large central perforations, smaller peripheral

perforations, margin smooth, up to 96 pm long, sometimes

partly bridged;

4. inner body wall, shallow bowls bridged on one side to create

smooth, hollow irregular ellipsoids, up to 88 pm long;

5. inner body wall, shallow bowls, frequently thick-walled,

bridged to create an upper surface and hollow ellipsoid, and

inner-bridged irregular ellipsoids, not hollow, typically up to

88 pm long, rarely up to 136 pm long, some becoming enlarged

and inter-grading with multi-layered scales;

6. underlying, multi-layered plates (scales), irregularly round

to oval, up to at least 1.6 mm across/long.

Tentacle ossicles (from NMV F210388) elongate, thick,

smooth, perforated rod-plates, curved and bent, up to 440 pm

long; fine distally perforate rods; rosettes, up to 96 pm long.

Tube feet ossicles (from NMV F210389) endplates,

uniform slightly irregular perforations, 400 /<m diameter;

endplate support rod-plates, narrow to elongate oval, to

rounded triangular, smooth or knobbed, curved and bent, up to

272 p m long; spinous-edge bowls, variably bridged, as in

dorsal body wall; shallow small bowls, variably knobbed, as in

dorsal body wall; shallow larger bowls, margin and surface

knobbed, two large and two smaller central perforations,

smaller peripheral perforations, up to 88 pm long; hollow and

inner-layered irregular ellipsoids as in dorsal body wall.

Live colour: radii and papillae variably red; dorsal and

lateral inter-radii variably greenish; ventral inter-radii pale

green; dendritic tentacles ends red; tentacle trunks greenish

yellow with dark brown to black flecking; ventral tube feet red.

Preserved colour: pale to dark grey.

Distribution. Through the tropical Indo-West-Pacific, from

Zanzibar to Malaysia and Australia; 0-115 m (depth from

Rowe & Gates 1995).

Remarks. We have observed specimens from Singapore waters

that we judge to be Colochirus quadrangular is. Because of the

proximal continuity of Singapore waters with those of the

Straits of Malacca, we judge that the Singapore specimens are

conspecific with those of the type locality. The live colour

photos of Colochirus quadrangularis from Singapore waters

that we have included here were provided for our work by our

colleagues Wong Pei San Helen and Joo Yong Ong (NUS

TMSI). The specimens studied here were donated to Museum

Victoria by the Lee Kong Chian Natural History Museum in

Singapore, the donation facilitated by our colleagues Wong Pei

San Helen and Joo Yong Ong. The description above

incorporates observations by these colleagues of 139 specimens

of this species in the Lee Kong Chian Natural History Museum.

The conspicuous and distinctive ossicle form in the upper

body wall of Colochirus quadrangularis is the irregular

shallow sub-rectangular bowl with one strongly spinous edge.

Leptopentacta grisea H. L. Clark, 1938

Table 1; appendix 1; figure 10a

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Linnaeus, sled, site no LIN_50,

barcode 53713, from -15.37072 124.69674 9 m to -15.36572 124.6978

9 m, 16 Mar 2015, WAM Z89027.

Plesiocolochirus Cherbonnier, 1946

Figure 1; appendices 1, 2

Acolochirus H. L. Clark, 1946: 395.—Panning, 1971: 41

(synonymy by Rowe in Rowe & Gates 1995)

Apentacta H. L. Clark, 1946: 395 (synonymy by Panning 1971,

and Clark & Rowe 1971)

Plesiocolochirus Cherbonnier, 1946: 286.—Panning, 1949: 448-

449.—Panning, 1971: 42.—Rowe (in Rowe & Gates), 1995: 277.

Type species. Plesiocolochirus spinosus (Quoy & Gaimard,

1834) (original designation)

Type species locality. Eastern Australia, New South Wales,

Port Jackson.

Other assigned species (with type locality added). Plesiocolochirus

armatus (Marenzeller von, 1881) (China); P. australis (Ludwig,

1875) (NE Australia, Bowen); P. challenged (Theel, 1886)

(N Australia, Torres Strait); P. dispar (Lampert, 1889)

(NW Australia, Mermaid Strait); P. ignavus (Ludwig, 1875) (S

Australia, Gulf St Vincent); P. inornatus (Marenzeller von, 1881)

(China); P. minaeus sp. nov. (NW Australia, Camden Sound,

below); P. minutus (Ludwig, 1875) (NE Australia, Bowen);

P. tessellarus (Cherbonnier, 1970) (Mozambique Channel).

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 10. a, photo of lateral view of preserved specimen of Leptopentacta grisea H. L. Clark, 1938 (WAM Z89027). b, c, photos of live

specimens of Plesiocolochirus species 1: b, dorsal view (WAM Z89028); c, lateral view (WAM Z89030).

P.M. O’Loughlin, C. Harding & G. Paulay

Diagnosis (sensu stricto, based on type species only,

described below). Body firm, packed with calcareous ossicles,

fusiform with upturned oral and anal ends, long posterior

taper, body oval in mid-body section, axial (not curved)

length up to 64 mm; lateral edges of body with prominent,

firm, conical papillae with imbricating scales; five conical

oral valves, with imbricating scales distally; anal cone with

large imbricating scales distally, large non-imbricating scales

anterior to imbricating ones; small anal scales; small ossicles

clustered into contiguous lumps in mid-body to create pseudo-

scales, each with central passage/perforation for withdrawn

tube feet, pseudo-scales about 1.00 mm across. Ten dendritic

tentacles, 8 large, 2 small. Calcareous ring cucumariid-like,

plates solid, higher than broad, lacking posterior prolongations.

Tube feet scattered irregularly over body, inconspicuous,

small, appear to penetrate pseudo-scales, more numerous

ventrally than dorsally, mid-ventral radius with irregular

paired rows, narrow space on each side of mid-ventral radius

lacking tube feet.

Three intergrading ossicle forms of body wall:

1. surface layer of finely knobbed, shallow bowls, variably

bridged across margins to sometimes create hollow, irregular,

ellipsoid-like ossicles;

2. layer of flat, thickly knobbed buttons, some developing

secondary layers to become small scales;

3. deeper layer of multi-layered scales.

Lateral papillae and tube feet with terminal endplates. Tentacle

ossicles rods; lacking rosettes.

Remarks. Cherbonnier (1946) proposed that the following

species be assigned to his new genus Plesiocolochirus:

Colochirus challengeri Theel, 1886; Colochirus gazellae

Lampert, 1889 (subsequently referred to Loisettea Rowe &

Pawson, 1985); Colochirus inornatus von Marenzeller, 1881;

Thyone papillata Sluiter, 1887 (subsequently referred to Stolus

Selenka, 1867); Colochirus squamatus Sluiter,

1901(subsequently synonymised with Colochirus robustus

Ostergren, 1898 by Rowe in Rowe & Gates 1995). Subsequently

other species have been assigned to Plesiocolochirus-.

Colochirus armatus Marenzeller von, 1881 (China); Colochirus

australis Ludwig, 1875 (NE Australia, Bowen); Colochirus

dispar Lampert, 1889 (NW Australia, Mermaid Strait);

Cucumaria ignava Ludwig, 1875 (S Australia, Gulf St Vincent);

Plesiocolochirus minaeus sp. nov. (NW Australia, Camden

Sound, below); Colochirus minutus Ludwig, 1875 (NE

Australia, Bowen); Pentacta tessellara Cherbonnier, 1970

(Mozambique Channel).

Rowe (in Rowe & Gates) 1995 referred Ocnus occiduus

O’Loughlin & O’Hara, 1992 to Plesiocolochirus, with

reservations. This species was subsequently referred to

Australocnus O’Loughlin & Alcock, 2000. In the same work

Rowe judged that Colochirus minutus is a junior synonym of

Plesiocolochirus australis. We provisionally raise Colochirus

minutus out of synonymy for further consideration.

Phylogenetic data based on COl sequences (Figure 1)

recover a clade of six species that appear to correspond to

Plesiocolochirus: P. challengeri (N Australia); P. ignavus

(SE Australia); P. minaeus sp. nov. (below) (NW Australia);

P. tessellarus (Comoros); Plesiocolochirus species 1 from

NW Australia and Palau; Plesiocolochirus species 2 from NE

Australia. The two unassigned species are close to specimens

that have frequently been referred to Plesiocolochirus

australis and Plesiocolochirus minutus, as well as to

Colochirus sp. 1 mentioned above. A detailed study of these

species, including study of relevant type specimens, is needed

to determine the identity of these species and morphological

boundaries of the Colochirus - Plesiocolochirus complex.

Unfortunately genetic data are not currently available for

Plesiocolochirus spinosus, the type species of the genus.

Morphologically this species is close to Plesiocolochirus

challengeri. We anticipate from morphological observations

that Plesiocolohirus spinosus, Plesiocolochirus challengeri,

Loisettea amphictena Rowe & Pawson, 1985 and Loisettea

gazellae will fall together in a clade distinct from the remaining

species assigned to Plesiocolochirus.

Plesiocolochirus Cherbonnier, 1946 (based on type species

Plesiocolochirus spinosus (Quoy & Gaimard, 1834), below) is

distinguished sensu stricto from Colochirus Troschel, 1846

(based on type species Colochirus quadrangularis Troschel,

1846, above) by the following characters:

1. presence of large imbricating scales on distal anal cone, on

proximal oral valves, and on lateral papillae;

2. presence of numerous inter-radial tube feet;

3. presence of knobbed button body wall ossicles;

4. absence of tentacle rosette ossicles.

Determination of the morphological limits of the clades

corresponding to these genera awaits further study.

Plesiocolochirus spinosus (Quoy & Gaimard, 1834)

Figures 11, 12

Holothuria spinosa Quoy & Gaimard, 1834: 118-120, pi. 7 figs

1 - 10 .

Cladolabes spinosus.— Brandt, 1835: 74.

Stolus firmus Selenka, 1867: 356, pi. 20 figs 118-119.

Ocnus spinosus.— Semper, 1867: 55.

Colochirus spinosus.— Selenka, 1868: 117—von Marenzeller,

1881: 129-132.—Theel, 1886,-Lampert, 1889: 825-826.

Thyone spinosa— Semper, 1869: 238.—Lampert, 1885: 157.

Stereoderma validum Bell, 1884: 150-151, pi. 9 figs Ea-f.

(synonymy by H. L. Clark 1946)

Apentacta spinosa.— H. L. Clark, 1946: 395.

Plesiocolochirus spinosus.— Cherbonnier, 1946: 280-286, fig.—

Rowe (in Rowe & Gates), 1995: 279.

Type locality. Australia, New South Wales, Port Jackson.

Material examined. S Queensland, Kimbla K4/69, -26.05 153.75 68 m,

1969, NMV F204081 (3); off Yeppoon, -24.05 151.45 9-37 m, 6 Sep

1967, NMV F204080 (1); off Yeppoon, Keppel Bay, -23.07 150.89 9

m, 6 Sep 1967, NMV F95257 (1).

Description. Body firm, packed with calcareous ossicles,

fusiform with upturned oral and anal ends, long posterior taper,

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

P.M. O’Loughlin, C. Harding & G. Paulay

Figure 12. Ossicles from specimen of Plesiocolochirus spinosus (Quoy & Gaimard, 1834) (NMV F204081): a, multi-layered scale ossicle from

body wall, up to 1.5 mm long; b, body wall thick buttons, up to 120 pm long, and fine knobbed bowls with (top center) and without (bottom left)

a bridge, up to 70 pm long; c, body wall bridged ellipsoid-like bowl (right), up to 70 pm long, and thick button (left).

body oval in mid-body section, dorsal length shorter than

ventral length, axial (not curved) length up to 64 mm; lateral

edges of body with 3-12 firm conical papillae with large

imbricating true scales, longest papillae mid-body, up to 6 mm

long; five conical oral valves, each with two prominent spines

distally, imbricating true scales distally; distal anal cone with

large imbricating true scales, interspersed with small ossicle

clumps; small anal scales; small ossicles clustered into

contiguous lumps in mid-body to create pseudo-scales, each

with central passage/perforation for withdrawn tube feet,

pseudo-scales about 1.00 mm across. Ten dendritic tentacles,

8 large, 2 small. Calcareous ring cucumariid-like, plates solid,

higher than broad, lacking posterior prolongations. Tube feet

scattered irregularly over body, frequently withdrawn and

inconspicuous, small, more numerous ventrally than dorsally,

mid-ventral radius with irregular paired rows, narrow space on

each side of mid-ventral radius lacking tube feet, tube feet

appear to penetrate pseudo-scales.

Three inter-grading ossicle forms of body wall:

1. surface layer of finely knobbed bowls, oval to rounded

rectangular in form, shallow concave, 4 central perforations,

variable number of additional comer perforations, finely knobbed

over surface, variably bridged from margins to sometimes create

hollow, irregular, ellipsoid-like ossicles, 40-70 pm long;

2. layer of thickly knobbed buttons, flat, large marginal and

central knobs, frequently 4 perforations but varying from 3-8,

buttons 50-120 pm long, some buttons developing secondary

layers to become small scales, up to about 216 pm long;

3. deeper layer of multi-layered scales, up to 1.5 mm long,

developed from additional layers on knobbed plates.

Lateral papillae and tube feet with terminal endplates, fairly

uniform perforations, about 120 pm.

Tentacle ossicles rods, lacking rosettes: large rods/rod-plates

thick, curved, bent, perforated along rod, variable form, up to

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

480 pm long; small, irregularly-branched rods, some

H-shaped, some distally perforate; fine very small distally

perforate rods, not branched.

Live colour (Cherbonnier 1946, based on Quoy & Gaimard

1834): red dorsally, grey ventrally, lateral spines purple,

tentacles red with brown spots at the base of the trunks.

Preserved colour (this work): off-white to pale brown with

some residual red flecks, two broad irregular transverse dark

brown bands around mid-body.

Distribution (Rowe & Gates 1995). Eastern Australia,

Queensland to Victoria, 9-90 m.

Plesiocolochirus species 1 (unresolved species complex)

Table 1; appendices 1, 2; figures 1, 10b, c

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander , sled, site no SOL_4,

WAM station no 22, barcode 10001662, from -15.446441 124.083021

62 mto -15.445955 124.083173 62 m, 20 Mar 2015, WAMZ89028 (1);

RV Solander , sled, site no SOL_24, WAM station no 23, barcode

10001822, from -15.406428 124.125928 42 m to -15.406937

124.125369 42 m, 20 Mar 2015, WAM Z89029 (1); RV Solander , sled,

site no SOL_43, WAM station no 26, barcode 10002101, from

-15.488461 124.201824 46 m to -15.488309 124.201113 46 m, 21 Mar

2015, WAM Z89030(l).

Remarks. We remarked under genus Plesiocolochirus (above)

that the phylogenetic tree (Figure 1) supports two discrete and

geographically separate Plesiocolochirus species clades from

NE Australia ( Plesiocolochirus species 2) and NW Australia

(Plesiocolochirus species 1). Either clade might be

representative of Plesiocolochirus australis ox Plesiocolochirus

minutus. Identification of the Camden species awaits

morphological examination of the relevant types and additional

phylogenetic data

Plesiocolochirus minaeus O’Loughlin sp. nov.

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act:

189703BE-67E5-4212-B337-8C5B2F39D1E8

Table 1; appendices 1, 2; figures 1, 13, 14;

Material examined. Holotype. Northwest Western Australia,

Kimberley Region, Camden Sound, WAMSI 1.1.1, RV Solander , sled,

site no SOL 117, WAM station no 31, barcode 10002268, from

-15.674833 124.279779 39 m to -15.674794 124.279012 39 m, 22 Mar

2015, WAM Z89026.

Description (preserved in 95% ethanol). Body hard, packed

with calcareous ossicles, elongate, square in transverse section,

70 mm long, 10 mm high and wide; surface of body wall

creased, imbricating scales around tips of papillae only; five

anterior oral valves, lobed terminal papillae on each valve;

each dorso-lateral margin with about 15 spaced, zig-zag,

pyramidal, hard papillae, up to 2 mm high; ventro-lateral

margin lacking papillae, except for 2-3 smaller papillae on

each of the three ventral oral radii, and one on each of the three

ventral anal radii; five tongue-like, radial anal scales/teeth; five

pyramidal, radial, anal papillae, not as high as dorso-lateral

papillae; dorsal and lateral inter-radii with scales evident,

irregular form and size, up to about 1.5 mm long. Ventral

surface flat with three broad, raised radii with lace-like network

of oblong scales each about 1 mm long, tube feet mostly deeply

retracted, ventro-lateral radial tube feet band about four wide,

mid-ventral band about five wide. Typical cucumariid

calcareous ring, undulating posteriorly, lacking posterior

prolongations. Ten dendritic tentacles, two ventral smaller.

Single polian vesicle, gonad tubules not branched.

Dorsal body wall and dorso-lateral papillae ossicles of six

intergrading types:

1. surface layer of deep bowls with tapering rounded base

and bluntly to sharply spinous or knobbed marginal rim,

some bridged internally, bowls typically about 55 pm wide

48 pm deep;

2. thick and thin walled shallow bowls, irregularly rectangular,

many with smooth rims, many with indented lateral rims,

variably bridged to create irregular hollow ellipsoids,

frequently 55 pm long, up to 144 pm long;

3. smooth, irregularly round to oval, hollow ellipsoids, up to

64 pm across;

4. some regular, four-holed, thickened, flat buttons, up to

72 pm long;

5. abundant knobbed and thickened irregular flat buttons,

many with incipient secondary layering, inter-grading with

small multi-layered scales, up to about 176 pm long;

6. multi-layered ossicles/scales, irregularly oval, up to at least

1.00 mm long.

Dorso-lateral papillae lacking apical tube feet and endplates.

Ventral tube feet ossicles of four forms (lacking multi-layered

ossicles and buttons):

1. endplates with fairly uniform perforations, smallest

centrally, endplates up to at least 280 pm diameter;

2. straight and curved, smooth, tube foot support rod-plates,

typically widened and perforated mid-rod and distally, some

marginally denticulate, rod-plates up to 200 pm long;

3. knobbed oval to rectangular shallow bowls, margins

knobbed to bluntly spinous, variably bridged to create irregular

hollow ellipsoids, bowls up to 55 pm long;

4. shallow bowls of variable size, not bridged, some with

marginal and surface knobs, bowls up to 128 pm long.

Tentacle ossicles of four inter-grading forms:

1. thick, smooth, perforated rod-plates up to 440 pm long;

2. smooth rods, variably perforated and branched

3. fine thin rods with distal perforations, typically about

60 pm long;

4. knobbed, branched rod rosettes, some perforated plates

with knobbed margin, up to 50 pm long.

P.M. O’Loughlin, C. Harding & G. Paulay

Figure 13. a, b, photos of live specimen of holotype of Plesiocolochirus minaeus O’Loughlin sp. nov. (WAM Z89026): a, dorsal view; b, ventral

view, c-e, photos of preserved holotype: c, lateral view; d, oral view; e, anal view.

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 14. Ossicles from holotype of Plesiocolochirus minaeus O’Loughlin sp. nov. (WAM Z89026): a, dorsal body wall deep bowls with

spinous rims about 55 jim wide 48 jim deep (bottom left and bottom center), shallow bowls variably bridged to create irregular hollow ellipsoids

up to 144 jim long (top row), knobbed button with secondary layering up to 176 jim long (bottom right); b, tentacle fine rod and thick perforated

rod up to 440 jim long, rosettes up to 50 ji m long; c, ventral tube foot endplate up to at least 280 jim long (right), support perforated rod-plates up

to 200 jim long, small shallow knobbed bowls variably bridged to create irregular hollow ellipsoids up to 55 jim long, large shallow bowl not

bridged up to 128 jim long (top left).

Live body colour very pale yellow to off-white, dorso-lateral

papillae and oral valves reddish-orange, ventral radii greenish

yellow; preserved colour off-white.

Distribution. Northwest Western Australia, Kimberley Region,

Camden Sound, 39 m.

Etymology. From the Latin minae (“parapets”), with reference

to the parapet-like hard papillae on the dorso-lateral margins of

the body in lateral view.

Remarks. The phylogenetic tree (Figure 1) includes a COl

sequence for the new species, Plesiocolochirus minaeus

O’Loughlin, within the congeneric clade of Plesiocolochirus

species. This sequence is understandably remote from the

Plesiocolochirus challengeri clade that we anticipate on

morphological grounds will be a clade close to

Plesiocolochirus spinosus.

The morphological characters that distinguish

Plesiocolochirus minaeus O’Loughlin sp. nov. from other

Plesiocolochirus species are the:

1. pyramidal, firm, dorso-lateral papillae;

2. complete absence of a ventro-lateral raised firm papillae;

3. absence of inter-radial tube feet;

4. presence of imbricating scales at tips of dorso-lateral

papillae only;

5. abundance of knobbed and thickened irregular buttons in

the body wall, many with incipient secondary layering, inter¬

grading with small multi-layered scales;

6. ventral tube feet surrounded by a ring of about four

ellipsoidal scales, not penetrating scales;

P.M. O’Loughlin, C. Harding & G. Paulay

7. live colour of red dorso-lateral papillae on off-white body.

We note the presence of tentacle rosettes in the new species.

This indicates that the presence or absence of tentacle rosettes

is not a sound generic diagnostic character for genera

Colochirus and Plesiocolochirus.

We also note the presence of two Pilumnidae crabs in the

coelom of the holotype of the new species.

Pseudocolochirus axiologus (H. L. Clark, 1914)

Table 1; appendix 1; figures 15, 16

Colochirus axiologus H. L. Clark, 1914: 171-173, pi. 25.—Ekman,

1918: 26-28, pi. 2 fig. 1, pi. 3 figs 16-19.

Pseudocolochirus axiologus (H. L. Clark, 1938): 456-457.—

1946: 394.

Pseudocolochirus violaceus (Theel, 1886).—Cherbonnier, 1988:

174-177, figs 73, 74 (part; N Australia specimens are P. axiologus).—

Rowe (in Rowe & Gates), 1995: 280 (part; N Australia specimens are

P. axiologus).

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no LIN_35,

WAM station no 23, barcode 10000760, from -15.363088 124.44389

37 m to -15.362756 124.443995 37 m, 17 Mar 2015, WAM Z89031 (1);

RV Solander, sled, site no SOL_49, WAM station no 32, barcode

10002389, from -15.668951 124.357909 41 m to -15.669007

124.357529 42 m, 22 Mar 2015, WAM Z89032 (1).

WA, northwest shelf, -12.84 125.68 88-97 m, 2 Apr 1981, NMV

F112187 (1); -12.90 125.58 84-88 m, 2 Feb 1981, NMV F112188 (2);

-12.90 125.59 83 m, 2 Apr 1981, NMV F112189 (2); -12.83 125.7 91 m,

1 Apr 1981, NMV FI 12190 (1).

Northern Territory, Joseph Bonaparte Gulf, -11.54 129.82 60 m,

12 Sep 2009, NMV F202986 (1) (tissue code MOL AF 1512); -10.31

129.62 89 m, 1 Sep 2009, NMV F202987 (1) (tissue code MOL AF

1504); -11.01 129.79 55 m„ 6 Sep 2009, NMV F202988 (2) (tissue code

MOL AF 1507).

Queensland, Gulf of Carpentaria, -11.37 141.42 35 m, 9 Sep 1982,

NMV F95259 (1) (tissue code MoV 4627).

Remarks. H. L. Clark (1914) noted the following features for

the type of his Colochirus axiologus:

1. 90 mm axial (horizontal) length;

2. tube feet confined to ventral ambulacra;

3. absence of ossicles in the body wall;

4. bright purple colour around the tentacle aperture;

5. body colour purplish-rose.

When describing additional specimens, H. L. Clark (1938)

referred his species to Pseudocolochirus Pearson, 1910, and

confirmed the absence of tube feet other than on the ventral

ambulacra, and the absence of ossicles in the body wall. H. L.

Clark (1938) acknowledged that Ekman (1918) found and

illustrated distinctive ossicles in the body wall of small

specimens (41-49 mm long) of what he judged to be Colochirus

axiologus , and Clark concluded that ossicles disappear with

increase in size of specimens.

We examined a small specimen (40 mm axial preserved

length) from Joseph Bonaparte Gulf (NMV F202988) that

lacked ossicles in the mid-body wall but did have a few almost

inconspicuous tube feet on the dorsal anterior radii (that H. L

Clark 1938 had also noted on his specimens). We also found

endplate support rod-plates in the ventral tube feet, and

distinctive small thick plates in the nearby body wall with very

small to no perforations. These ossicles were frequently

dumbbell-shaped and lacked perforations, or had up to three

very small ones. The plates varied in size from 40-90 pm

long. We found similar small plates near the ventral tube feet

in a larger specimen (100 mm axial preserved length) from

Camden Sound (WAM Z89032). These buttons were larger, up

to 200 pm long, and more irregular in form. There were a few

anterior dorsal radial hard papillae in which we found multi¬

layered ossicles fragments, and large single-layered perforated

plate fragments up to 440 pm long.

We judge that the specimens examined by H. L. Clark

(1914, 1938), Ekman (1918) and us are conspecific and belong

to Pseudocolochirus axiologus (H. L. Clark, 1914). We

acknowledge that this species is similar to Pseudocolochirus

violaceus (Theel, 1886). We do not accept the synonymy of

these two species by Cherbonnier (1988). Pseudocolochirus

violaceus has the following differing characters:

1. the whole body, both radial and inter-radial, is covered with

small papillae;

2. tube feet are clearly evident on the dorsal radii;

3. the distinctive plates are present in the mid-body wall;

4. prominent anterior and posterior papillae are more

numerous.

We raise Pseudocolochrus axiologus (H. L. Clark, 1914) out

of synonymy (by Cherbonnier 1988) with Pseudocolochirus

violaceus (Theel, 1886). Rowe (in Rowe & Gates, 1995)

followed Cherbonnier (1988) who considered all

Pseudocolochirus species to be synonyms, with

Pseudocolochirus violaceus the senior synonym. We judge

that northern Australian specimens are Pseudocolochirus

axiologus, not P. violaceus (see synonymy above).

Family Phyllophoridae Ostergren, 1907 ( sensu Pawson & Fell

1965)

Phyllophorella spiculata (Chang, 1935)

Table 1; appendix 1; figure 17b

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no SOL_32,

WAM station no 19, barcode 10001306, from -15.253592 124.203038

45 m to -15.253318 124.202302 45 m, 19 Mar 2015, WAM Z89033 (1).

Remarks. We note that O’Loughlin et al. (2012) raised

Phyllophorella to generic status.

Phyllophorus (Urodemella) holothurioides Ludwig, 1875

Table 1; appendix 1; figure 17a

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 15. Photos of live specimen of Pseudocolochirus axiologus (H. L. Clark, 1914) (WAM Z89032): a, lateral view; b, peri-oral view; c, peri¬

anal view.

P.M. O’Loughlin, C. Harding & G. Paulay

Figure 16. Ossicles from specimen of Pseudocolochirus axiologus (H. L. Clark, 1914) (WAM Z89031). From ventral tube feet and ventral body

wall near tube feet: thick irregular oval to elongate plates with small perforations, plates up to 200 pim long; tube foot thick and thin support

plates with distal small perforations, plates up to 256 jtm long; small endplate (mid-lower).

Material examined. Northwest Western Australia, Kimberley

Region, Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no

LIN_46, WAM station no 3, barcode 10000113, from -15.399083

124.345337 41 m to -15.398828 124.345572 42 m, 15 Mar 2015,

WAM Z89034 (1); RV Solander, sled, site no SOL_116a, WAM

station no 18, barcode 10001277, from -15.261423 124.275183 41 m

to -15.261716 124.275827 41 m, 18 Mar 2015, WAM Z89035 (1).

Family Sclerodactylidae Panning, 1949 ( sensu Smirnov 2012)

Havelockia Pearson, 1903

Havelockia Pearson, 1903: 198.—Panning, 1949: 466, 468.—

Rowe (in Rowe & Gates), 1995: 310.

Pentathyone Clark, H. L„ 1938: 458-459.-1946: 386, 396.-

Panning, 1949: 459.

Diagnosis (after Thandar 1989). Calcareous ring short, stout,

only anterior projections of radial and inter-radial plates free;

posterior paired process of radial plates divided into several

pieces. Body wall ossicles tables with squarish to oval discs,

usually perforated by four large central and four smaller

peripheral perforations, that are sometimes reduced or absent;

spire of two pillars joined at apex and terminating in a few

blunt teeth.

Remarks. Thandar (1989) has discussed the above generic

synonymy by Panning (1949), and the replacement of the type

species Havelockia herdmani Pearson, 1903 by the senior

synonym Havelockia versicolor (Semper, 1867).

Havelockia versicolor (Semper, 1867)

Table 1; appendix 1; figures 18a-d

Material examined. Northwest Western Australia, Kimberley

Region, Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no

SOL_120, WAM station no 5, barcode 10000277, from -15.37669

124.248319 53 m to -15.377306 124.248179 53 m, 15 Mar 2015,

WAM Z89036 (1); RV Solander, sled, site no LIN_35, WAM station

no 11, barcode 10000776, from -15.363088 124.44389 37 m to

-15.362756 124.443995 37 m, 17 Mar 2015, WAM Z89037 (1); RV

Solander, sled, site no LIN_6, WAM station no 16, barcode 10001220,

from -15.27505 124.36988 46 m to -15.27543 124.369177 46 m, 18

Mar 2015, WAM Z89038 (1); RV Solander, sled, site no LIN_36,

WAM station no 17, barcode 10001124, from -15.220444 124.320894

50 m to -15.220159 124.320648 50 m, 18 Mar 2015, WAM Z89039

(1); RV Solander, sled, site no SOL_32, WAM station no 19, barcode

10001321, from -15.253592 124.203038 45 m to -15.253318

124.202302 45 m, 19 Mar 2015, WAM Z89040 (1); RV Solander,

sled, site no SOL_56, WAM station no 20, barcode 10001417, from

-15.376537 124.192773 35 m to -15.376196 124.192071 35 m, 19 Mar

2015, WAM Z89041 (1); RV Solander, sled, site no SOL_4, WAM

station no 22, barcode 10001637, from -15.446441 124.083021 62 m

to -15.445944 124.083173 62 m, 20 Mar 2015, WAM Z89042 (1); RV

Solander, sled, site no SOL_24, WAM station no 23, barcode

10001924, from -15.406428 124.125928 42 m to -15.406937

124.125369 42 m, 20 Mar 2015, WAM Z89043 (1); RV Solander, sled,

site no SOL_49, WAM station no 32, barcode 10002314, from

-15.668951 124.357909 41 m to -15.669007 124.357529 42 m, 22 Mar

2015, WAM Z89044 (1); RV Solander, sled, site no SOL_49, WAM

station no 32, barcode 10002323, from -15.668951 124.357909 41 m

to -15.669007 124.357529 42 m, 22 Mar 2015, WAM Z89045 (1); RV

Solander, sled, site no SOL_73, WAM station no 38, barcode

10002688, from -15.945442 124.366373 29 m to -15.945268

124.367171 28 m, 25 Mar 2015, WAM Z89046 (1); RV Solander, sled,

site no SOL_73, WAM station no 38, barcode 10002689, from

-15.945442 124.366373 29 m to -15.945268 124.367171 28 m, 25 Mar

2015, WAM Z89047 (1); RV Solander, sled, site no SOL_97, WAM

station no 39, barcode 10002752, from -15.782865 124.378047 32 m

to -15.782335 124.378553 33 m, 25 Mar 2015, WAM Z89048 (1); RV

Solander, sled, site no SOL_69, WAM station no 41, barcode

10002838, from -15.747648 124.146502 43 m to -15.747285 124.14634

43 m, 26 Mar 2015, WAM Z89049 (1); RV Solander, sled, site no

SOL_47, WAM station no 42, barcode 10002939, from -15.612805

124.073033 3 6 m to -15.612437 124.072883 35 m, 26 Mar 2015,

WAM Z89050 (1).

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 17. a, photo of dorsal view of live specimen of Phyllophorus (Urodemella) holothuroides Ludwig, 1875 (WAM Z89035). b, photo of

lateral view of live specimen of Phyllophorella spiculata (Chang, 1835) (WAM Z89033).

Family Thyonidae Panning, 1949 ( sensu Smirnov, 2012)

Subfamily Semperiellinae Heding & Panning, 1954

Diagnosis. See O’Loughlin, Mackenzie & VandenSpiegel

2014.

Remarks. See O’Loughlin, Mackenzie & VandenSpiegel 2014.

Massinium Samyn & Thandar, 2003

Massinium Samyn and Thandar, 2003: 136.—Samyn et al., 2010:

2.—O’Loughlin et al., 2012: 290.

Diagnosis (O’Loughlin, Mackenzie & VandenSpiegel 2014).

Frequently semi-spherical species with oral and anal dorsal

orientations; 20 dendritic tentacles arranged in two circles of

10 large outer and 10 small inner (proximal peri-oral); tube feet

distributed all over mid-body; calcareous ring elongate, tubular,

with both radial and inter-radial plates fragmented into a

mosaic of small pieces, and posterior prolongations linked

distally to form inter-radial oval non-calcified spaces beneath

the water vascular ring; polian vesicles from 1 to 4; ossicles

variably include granuliform rods, rosettes, pseudo-buttons

and tables; table spires with 1 or 2 or 3 or reduced pillars.

Remarks. See O’Loughlin, Mackenzie & VandenSpiegel 2014.

Massinium bonapartum O’Loughlin, 2014 (in O’Loughlin,

Mackenzie & VandenSpiegel 2014)

Table 1; appendix 1; figure 19d

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander , sled, site no SOL_2,

WAM station no 30, barcode 10002243, from -15.711306 124.315779

28 m to -15.71207 124.31566 28 m, 22 Mar 2015, WAM Z89051 (1);

RV Solander , sled, site no SOL_73, WAM station no 38, barcode

10002725, from -15.945442 124.366373 29 m to -15.945268

124.367171 28 m, 25 Mar 2015, WAM Z89052 (1); RV Solander , sled,

site no SOL_47, WAM station no 42, barcode 10002966, from

-15.612805 124.073033 36 m to -15.612437 124.072883 35 m, 26 Mar

2015, WAM Z89053 (1).

P.M. O’Loughlin, C. Harding & G. Paulay

Figure 18. Photos of live specimens of Havelockia versicolor (Semper, 1867) (estimated 35-40 mm long live), a, dorsal view (WAM Z89047); b,

ventral view (WAM Z89047); in situ view (WAM Z89043); d, lateral view (WAM Z89048).

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Neothyonidium (?) insolitum O’Loughlin sp. nov.

Zoobank LSID. http://zoobank.Org/urn:lsid:zoobank.org:act:

7FCA7216-B862-4A4E-A829-07CA9DC0E23D

Table 1; appendix 1; figures 19a-c, 20

Material examined. Holotype. Northwest Western Australia,

Kimberley Region, Camden Sound, WAMSI 1.1.1, RV Solander, sled,

site no SOL_47, WAM station no 42, barcode 10002958, from

-15.612805 124.073033 36 m to -15.612437 124.072883 35 m, 26 Mar

2015, WAM Z89054.

Description. Body fusiform, cylindrical mid-body, tapered and

rounded oral end, tapered anally to narrow end, live body

variably 50 mm long, 13 mm diameter, body (preserved in 95%

ethanol) 26 mm long, up to 14 mm diameter; complete cover of

tube feet, more numerous ventrally than dorsally, tube feet

diameters about 0.2 mm; tentacles in inner circle of five very

small, outer circle of 13 large; calcareous ring elongate, tubular,

with both radial and inter-radial plates fragmented into a

mosaic of small pieces; inconspicuous anal scales detected;

single polian vesicle.

Ossicles of mid-body wall numerous Thyone-Wke oval

tables, discs with most frequently four perforations, sometimes

with an additional four smaller corner perforations to create a

rounded rectangular disc, discs up to 80 pm long; spires two

short pillars with one distal cross-bridge, two distal splayed

spines at base of each pillar, spire height about 24 pm. Tube

feet endplates with variable diameters up to 224 pm, irregular

perforations and margin; rare tube foot support tables with

elongate discs up to 80 pm long; some tube foot support rods,

similar to tentacle rods, sometimes forked distally with

enlarged rounded ends with few small perforations. Peri-oral

wall with tables similar to mid-body wall, but discs generally

smaller, up to 55 pm long. Peri-anal body wall with tables

similar to mid-body wall, but smaller as in peri-oral region;

small rosettes present; anal multi-layered scales present.

Tentacles with thick and thin rods, distal ends swollen with

small perforations; thick rods sometimes branched distally

into widened ends with small perforations, thick rods up to

160 pm long; thin rods up to 90 pm long.

Live colour off-white to grey, semi-translucent; preserved

colour off-white.

Distribution. Northwest Western Australia, Kimberley Region,

Camden Sound, 35-36 m.

Etymology. From the Latin insolitum (meaning unusual), with

reference to the unusual combination of characters for the

genus Neothyonidium.

Remarks. We refer the new species to Neothyonidium

Deichmann, 1938 on the basis of the body form, cover of tube

feet, near 20 tentacles of two sizes, form of the composite

calcareous ring, and body wall table ossicles with two pillars.

But we do so with reservation because of the atypical

combination of two morphological characters: five very small

tentacles and 13 larger ones; Thyone-\ike table ossicles in the

body wall. These two characters distinguish Neothyonidium(2)

insolitum O’Loughlin from all other Neothyonidium species.

Subfamily Thyoninae Panning, 1949

Hemithyone semperi (Bell, 1884)

Table 1; appendix 1; figure 21

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no SOL_56,

WAM station no 20, barcode 10001531, from -15.376537 124.192773

35 m to -15.376196 124.192071 35 m, 19 Mar 2015, WAM Z89055 (1).

Remarks. We have retained Hemithyone semperi (Bell, 1884) in

family Thyonidae and subfamily Thyoninae, but with major

reservations. The species does have a composite calcareous ring,

but the tube feet are radial and the species lacks table and cup

ossicles and has predominantly very open fenestrated ellipsoids

in the body wall. Smirnov (2012) remarked “it is possible that

Hemithyone Pawson, 1963 does not belong to Thyonidae”. We

await generic evidence for a review of its higher taxon referral.

Stolus canescens (Semper, 1867)

Table 1; appendix 1; figure 22

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no SOL_2,

WAM station no 30, barcode 10002195, from -15.711306 124.315779

28 m to -15.71207 124.31566 28 m, 22 Mar 2015, WAM Z89056 (1).

Thy one Oken, 1815

Diagnosis (emended in O’Loughlin et al. 2012 from Pawson

and Miller 1981). Tentacles 10; tube feet scattered on body

wall, never restricted to radii; calcareous ring tubular with long

posterior prolongations comprising a mosaic of small pieces;

body wall ossicles tables with a spire of one or two pillars.

Type species. Holothuria fusus O. F. Muller, 1776 (monotypy).

Northern Australia species q/Thyone reported in Rowe &

Gates 1995 (type locality added). T. axiologa H. L. Clark,

1938 (Broome); T. dura Koehler & Vaney, 1908 (W India)

(junior synonym T. alba H. L. Clark, 1938, by Heding 1940

(Broome)); T. grisea H. L. Clark, 1938 (Cape Bossut, N

Australia); T. micra H. L. Clark, 1938 (Broome); T. papuensis

Theel, 1886 (Torres Strait).

Remarks. We note in the Introduction the recent ruling by

the ICZN that Thyone Oken, 1815 is now an available

taxon. Pawson & Miller (1981) remarked on the need for a

revision of the “supergenus” Thyone. Arumugam (2012) has

provided a morphological approach to the “management” of

this “supergenus”.

Liao & Clark (1995) noted that the holotype specimen of

Thyone papuensis is now very damaged and completely

decalcified. The original description and illustrations of the

species are not sufficient for diagnostic comparisons and we

thus provide (below) a description of specimens that we judge

to be Thyone papuensis.

We add Thyone pedata Semper, 1867 to northern Australia

species of Thyone on the basis of a specimen identified by us

from Joseph Bonaparte Gulf (NMV F173267; UF tissue lot

MOL AF 1537). We note that Rowe (in Rowe & Gates 1995)

P.M. O’Loughlin, C. Harding & G. Paulay

Figure 19. a-c, photos of holotype of Neopsolidium(l) insolitum O’Loughlin sp. nov. (WAM Z89054): a, photo of dorsal view of live holotype;

b, photo of lateral view of preserved holotype; c, photo of calcareous ring of preserved holotype. d, photo of live specimen of Massinium

bonapartum O’Loughlin, 2014 (in O’Loughlin, Mackenzie & VandenSpiegel, 2014) (WAM Z89051; estimated 20 mm long live).

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 20. Ossicles from holotype of Neopsolidiumil ) insolitum O’Loughlin sp. nov. (WAM Z89054): a, from tentacle, thick rods with widened

and branched perforate ends up to 160 pm long, thin rods up to 90 pm long; b, from dorsal body wall, table discs up to 80 pm long, low two-pillar

spires, distinctive tube foot support rod with bifurcate perforate ends; c, from posterior body wall, multi-layered scale fragment (bottom left),

endplate up to 224 pm diameter (bottom right), distally widened and perforate tube foot support rods (top right), table discs up to 55 pm long

(center), rosettes (top left).

referred Thyone perissa H. L. Clark, 1938 (WA) to Massinium

magnum (Ludwig, 1882), and Thyone minuta H. L. Clark,

1938 to Stolus minutus (H. L. Clark, 1938).

Thyone papuensis Theel, 1886

Table 1; appendix 1; figures 23, 24

Thyone fusus var. papuensis Theel, 1886: 92, pi. 7 fig. 1.

Thyone papuensis H. L. Clark, 1921: 167.-1932: 221.-1946:

399.—Clark & Rowe, 1971: 182.-A. M. Clark, 1982: 489, 495, fig.

2.—Cannon & Silver, 1986: 32, fig. 9g.—Liao & Clark, 1995: 504, fig.

306.—Rowe (in Rowe & Gates), 1995: 316.

Type locality. Torres Strait.

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no LIN_6,

WAM station no 16, barcode 10001097, from -15.27505 S 124.36988

E 46 m to -15.27543 S 124.36918 E 46 m, 18 Mar 2015, WAM Z89057

(1); RV Solander, sled, site no SOL_116a, WAM station no 18, barcode

10001259, from -15.26142 S 124.27518 E 41 m to -15.26172 S

124.27583 E 40 m, 18 Mar 2015, WAM Z89058 (1); RV Solander, sled,

site no SOL_43, WAM station no 26, barcode 10002102, from

-15.488461 124.201824 46 m to -15.488309 124.201113 46 m, 21 Mar

2015, WAM Z89059 (1) (ring eviscerated and lost); RV Solander, sled,

site no SOL_77, WAM station no 34, barcode 10002542, from

-15.725854 S 124.166978 41 m to -15.726405 124.16756 41 m, 23 Mar

2015, WAM Z89060(l).

North Kimberley Region, near mouth of King George River, RV

Solander, Sled 06, lot number (barcode) 023319, -13.8505 127.28868

45 m, 6 June 2013, WAM Z27863 (1) (UF tissue lot MOL AF 1463).

Description (Kimberley specimens; preserved in 95% ethanol).

Body fusiform, narrow, with long tapers to narrow oral and

anal ends, oral and anal ends may be slightly upturned, body

(live) up to 35 mm long, up to 17 mm diameter; complete cover

P.M. O’Loughlin, C. Harding & G. Paulay

Figure 21. Photo of live specimen of Hemithyone semperi (Bell, 1884) (WAM Z89055).

of small tube feet, diameters about 0.2 mm, some in longitudinal

series; 10 dendritic tentacles, two ventral small; calcareous ring

tubular with long posterior prolongations comprising a mosaic

of small pieces; inconspicuous anal scales present comprising

thick single-layered plates with small perforation, multi-layered

components, meshed rods component; single polian vesicle;

gonad tubules not branched.

Mid-body ossicles scattered small tables with a spire of

two pillars, discs with predominantly four perforations in

smaller specimens, two larger central, two smaller distal,

many discs in larger specimens with four or more smaller

additional perforations, disc lengths commonly 50-55 pm,

spire heights 23 pm, spires of two pillars, single apical bridge,

apex of spire with two short pillar ends, each with a few blunt

spines, spines sometimes with additional secondary spinelet.

Tube feet endplates Thyone-Yike, with small central

perforations, outer ring of large perforations, marginal rim of

small transversely oval perforation, endplates up to 110 pm

diameter; support tables in tube feet with elongate, curved

discs up to about 120 pm long, four central perforations, single

small perforation distally; spires with two pillars, joined

apically with few blunt apical spines. Peri-anal body wall with

single and multi-layered scale ossicles, tables with multi-

perforate discs, tube foot support tables differing from mid¬

body. Tentacles and introvert with fine rods, tables and

rosettes; largest rods with bifurcate ends, rods up to 90 pm

long; table discs with many perforations, up to about 20;

rosettes typically oval, about 30 pm long; rods and rosettes

inter-grade in form.

Live colour pale brown to yellow with irregular red to

brown patches on body and some tube feet; preserved colour

off-white with red-brown to brown patches on body and some

tube feet.

Distribution. North Western Australia, Kimberley Region,

Camden Sound and King George River region, 40-46 m (this

work); Houtman Abrolhos, WA, to Double Island Point,

Queensland, 0-60 m (Rowe & Gates 1995).

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 22. Photo of live specimen of Stolus canescens (Semper, 1867) (WAM Z89056).

P.M. O’Loughlin, C. Harding & G. Paulay

Figure 23. Photos of specimens judged to be Thyone papuensis Theel, 1886. a, lateral view of live specimen (WAM Z89060); b, calcareous ring

of the same specimen preserved; c, lateral view of live specimen (WAM Z89058); d, calcareous ring of same specimen preserved (ring is

relatively very small and is probably regenerating).

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Figure 24. Photos of ossicles from Thyone papuensis Theel, 1886 specimens (WAM Z89057, WAM Z89058, WAM Z89059). a, from tentacle

and introvert, fine rods (up to 90 pm long), rosettes (frequently about 30 pm long), table with multi-perforate disc and two spires; b, from mid¬

body wall and tube foot, table discs (frequently 50-55 pm long) and spires (about 23 pm high), endplate (up to 110 pm diameter), endplate support

tables with single distal disc perforations (curved discs up to about 120 pm long); c, from peri-anal body wall, table with multi-perforate disc 56

pm across, tube foot support tables differing from body wall, upper one 80 pm between distal ends, lower left one 120 pm long; d, from peri-anal

body wall, single and multi-layered fragments of scale ossicle.

Family Thyonidiidae Heding & Panning, 1954 ( sensu Smirnov

2012 )

Remarks. See O’Loughlin, Mackenzie & VandenSpiegel, 2014.

Actinocucumis Ludwig, 1875

Remarks. See O’Loughlin, Mackenzie & VandenSpiegel, 2014.

Actinocucumis longipedes H. L. Clark, 1938

Table 1; appendix 1; figure 25a

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no SOL_40,

WAM station no 10, barcode 10000683, from -15.339415 124.160692

46 m to -15.33941 124.161459 46 m, 16 Mar 2015, WAM Z89061 (1).

Actinocucumis solanderi O’Loughlin, 2014 (in O’Loughlin,

Mackenzie & VandenSpiegel, 2014)

Table 1; appendix 1; figure 25b

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no LIN_46,

WAM station no 3, barcode 10000111, from -15.399083 124.345337

41 m to -15.398828 124.345572 42 m, 15 Mar 2015, WAMZ89062 (1);

RV Solander, sled, site no LIN_36, WAM station no 17, barcode

10001112, from -15.220444 124.320894 50 m to -15.220159

124.320648 50 m, 18 Mar 2015, WAM Z89063 (1); RV Solander, sled,

site no SOL_69, WAM station no 41, barcode 10002832, from

-15.747648 124.146502 43 m to -15.747285 124.14634 43 m, 26 Mar

2015, WAM Z89064 (1).

P.M. O’Loughlin, C. Harding & G. Paulay

Figure 25. Photos of lateral views of live specimens of three species of Actinocucumis Ludwig, 1875: a, Actinocucumis longipedes H. L. Clark,

1938 (WAM Z89061); b, Actinocucumis solanderi O’Loughlin, 2014 (in O’Loughlin, Mackenzie & VandenSpiegel, 2014) (WAM Z89063); c,

Actinocucumis typica Ludwig, 1875 (WAM Z89065; estimated 25 mm long live).

Actinocucumis typica Ludwig, 1875

Table 1; appendix 1; figure 25c

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no SOL_101,

WAM station no 35, barcode 10002613, from -15.753636 124.270741

34 m to -15.75404 124.271287 34 m, 23 Mar 2015, WAM Z89065 (1);

RV Solander, sled, site no SOL_97, WAM station no 39, barcode

10002792, from -15.782865 124.378047 32 m to -15.782335

124.378553 33 m, 25 Mar 2015, WAM Z89066 (1).

Mensamaria intercedens (Lampert, 1885)

Table 1; appendix 1; figure 26

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no SOL_40,

WAM station no 10, barcode 10000685, from -15.339415 124.160692

46 m to -15.33941 124.161459 46 m, 16 Mar 2015, WAM Z89067 (1);

RV Solander, sled, site no SOL_32, WAM station no 19, barcode

10001307, from -15.253592 124.203038 45 m to -15.253318

124.202302 45 m, 19 Mar 2015, WAM Z89068 (1).

Order Molpadida Haeckel, 1896

Family Molpadiidae J. Muller, 1850

Molpadia scabrum (Sluiter, 1901)

Table 1; appendix 1; figure 27a

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Linnaeus, sled, site no LIN_48,

barcode 53720, from-15.32474 124.42657 28 m to-15.32183 124.42742

12 m, 19 Mar 2015, WAM Z89069 (1).

Subclass Synaptacea Cuenot, 1891 {sensu Smirnov 2012)

Order Synaptida Cuenot, 1891 {sensu Smirnov 2012)

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Family Synaptidae Burmeister, 1837 {sensu Ostergren 1898)

Subfamily Rynkatorpinae Smirnov, 1989

Protankyra Ostergren, 1898

Diagnosis (after Clark 1908). Tentacles digitate, 10-12, rarely

13 or 14; digits two on each side (rarely one only). Cartilaginous

ring wanting. Polian vesicles 2-10, rarely one only. Stone canal

usually single, rarely several. Stock of anchors more or less

branched or only finely toothed; arms usually serrate; vertex of

anchors without knobs. Anchor plates without a handle; with

numerous irregular perforations, never with two large central

perforations; with a more or less imperfectly developed bow

across outer surface of posterior end; plates and perforations

with either smooth or dentate margins.

Remarks. We have added “never with two large central

perforations” to the diagnosis of anchor plates to distinguish

Protankyra from genus Rynkatorpa Rowe & Pawson, 1967.

Protankyra insolens (Theel, 1886)

Table 1; appendix 1; figure 27b

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no SOL_107,

WAM station no 1, barcode 10000038, from-15.514826 124.183111 46

m to -15.514503 124.183774 45 m, 14 Mar 2015, WAM Z89070 (1).

Protankyra torquea O’Loughlin sp. nov.

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act:

94FD3258-A AE8-48F3 -939F-444D8ECC70FD

Table 1; appendix 1; figure 28

Material examined. Holotype. Northwest Western Australia,

Kimberley Region, Camden Sound, WAMSI 1.1.1, RV Solander , sled,

site no SOL_48, WAM station no 4, barcode 10000142, from -15.39822

124.28823 23.2 m to -15.39864 124.28950 28.6 m, 15 Mar 2015, WAM

Z89071.

Description (preserved in 95% ethanol). Anterior end only of

synaptid species, 26 mm long, up to 16 mm diameter; body

wall firm to hard, thick, opaque, not translucent; tentacles 12,

trunks elongate, each with two pairs of closely placed distal

digits, distal end of tentacles with short, thick papilla-like end;

four polian vesicles detected; ciliated funnels not detected at

base of dorsal mesentery or along coelomic inter-radii;

longitudinal muscles broad, thick, rounded, undivided.

Body wall ossicles anchors, anchor plates, short rods,

miliary granules. Anchors irregular in form, similar sizes, up

to 304 pm long, base of stocks variably indented, variably

finely toothed, lateral ends of stock with raised rounded

elevations, shaft variably constricted near base, anchor vertex

lacking knobs, arms with 3-5 outer anteriorly pointed spines.

Anchor plates irregularly heart-shaped, breadth and height

sub-equal, size slightly variable, plates 230-260 pm long,

perforations (excluding basally) up to about 30, irregular in

size, basal perforations numerous and small, fine teeth on

inner margin of perforations numerous to rare to absent, plates

lacking significantly larger central perforations, margin of

plates incomplete and irregular, irregular bow across posterior

end of plates. Short curved rods abundant, of variable form,

some curved inwards distally, some with distal swellings,

some bluntly denticulate on inner margin, rods up to about 30

pm long. Some miliary ‘granules’ found in body wall, not

abundant, thin oval flat plates, some dumbbell shaped, up to

about 30 pm long. Tentacles with miliary ‘granules’ and rods,

as in body wall. Longitudinal muscles with abundant miliary

‘granules’ only, ‘granules’ as in body wall and tentacles.

Live body colour off-white with irregular red-brown

transverse patches, tentacle trunks as for body, digits pale

yellow; preserved body colour off-white with irregular pale

brown patches.

Distribution. Northwest Western Australia, Kimberley Region,

Camden Sound, 23-29 m.

Etymology. Named torquea from the Latin torqueo (irregular),

with reference to the irregular form of the typically incomplete

margin of anchor plates, and irregular form of the anchor stock

bases.

Remarks. The specimen comprises an anterior end only, is

strongly contracted, and the alimentary canal and mesentery

are mostly eviscerated. These factors might account for the

apparent absence of ciliated funnels. Three Protankyra species

are reported from northern Australia by Rowe (in Rowe &

Gates 1995): P. insolens (Theel, 1886) (type locality Arafura

Sea, north of Camden Sound); P. similis (Semper, 1867) (type

locality the Philippines); P. verrilli (Theel, 1886) (type locality

Torres Strait, NE Australia). Both P. insolens and P. verrilli

were found in Camden Sound and are reported here. Each fits

well with the description and illustration by Theel (1886).

Amongst Protankyra species, Protankyra torquea O’Loughlin

sp. nov. is closest to P. verrilli in morphological characters, but

P. verrilli lacks rods in the body wall, is smaller, has a thin

body wall, and lacks colour in the preserved state.

Protankyra verrilli (Theel, 1886)

Table 1; appendix 1; figure 27c

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no SOL_47,

WAM station no 42, barcode 10002967, from -15.612805 124.073033

36 m to -15.612437 124.072883 35 m, 26 Mar 2015, WAM Z89072 (1).

Subfamily Synaptinae Burmeister, 1837 ( sensu Smirnov 1989)

Synaptula lamperti Heding, 1928

Table 1; appendix 1; figure 29a

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no SOL_56,

WAM station no 20, barcode 10001534, from -15.376537 124.192773

35 m to -15.376196 124.192071 35 m, 19 Mar 2015, WAM Z89073 (1).

Synaptula recta (Semper, 1867)

Table 1; appendix 1; figure 29b

P.M. O’Loughlin, C. Harding & G. Paulay

Figure 26. Photo of live specimen of Mensamaria intercedens (Lampert, 1885) (WAM Z89067).

Material examined. Northwest Western Australia, Kimberley Region,

Camden Sound, WAMSI 1.1.1, RV Solander, sled, site no LIN_35,

WAM station no 11, barcode 10000777, from -15.363088 124.443893

37 m to -15.362756 124.443995 37 m, 17 Mar 2015, WAM Z89074 (1);

same data, barcode 10000919, WAM Z89075 (1).

Acknowledgements

We are grateful to the following for their invaluable assistance

with our paper: John Keesing (CSIRO) for his collaborative

assistance with loans, data and photos; Joo Yong Ong and

Wong Pei San Helen (NUS, TMSI) for facilitating the

provision of live colour specimen photos and the donation of

Singapore specimens; the Lee Kong Chian Natural History

Museum in Singapore for the donation of specimens; Chris

Mah (USNM) for assistance with literature; Melanie

Mackenzie (NMV) for assistance with photography; Frangois

Michonneau (UF) for confirming a species identity; Kate

Naughton (NMV) for facilitating loan transport; Frank Rowe

(Research Associate of the Australian Museum) for his

valuable opinions on some of the systematics; Ken Walker

(NMV) for facilitating our use of the NMV Entomology

Department microscopes; and Western Australian Museum

staff members for assistance with data, literature and loans

(Clay Bryce, Jane Fromont, Loisette Marsh, Jenelle Ritchie,

Stacey Osborne, and Mark Salotti). We are most appreciative

of the contribution of Alex Kerr (University of Guam) for his

critical review of our paper.

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Samyn, Y., Thandar, A. S. & VandenSpiegel, D. 2010. Two new species

in the phyllophorid genus Massinium (Echinodermata:

Holothuroidea) with redescription of Massinium magnum. Zootaxa

2399: 1-19.

Selenka, E. 1867. Beitrage zur Anatomie und Systematik der Holothurien.

ZeitschriftJur Wissenschaftliche Zoologie 17(2): 291-374, pis 17-20.

Selenka, E. 1868. Nachtrag zu den Beitragen zur Anatomie und

Systematik der Holothurien. Zeitschrift Wissenschaftliche

Zoologische 18: 109-119, pi. 8.

Semper, C. 1867 (1868). Reisen im Archipel der Philippines Zweiter

Theil. Wissenschaftliche Resultate. 1, Holothurien. 285 pp, 40 pis.

Wilhelm Engelmann, Leipzig.

Semper, C. 1869. Die Holothurien Ostafrika’s. Baron Carl Claus von der

Decken’s Reisen in Ost-Afrika in den Jahren 1859-1865.

Wissenschaftlicher Theil 3 (1): 117-122.

Sluiter, C. P. 1887. Die Evertebraten aus der Sammlung des Koniglichen

Naturwissenschaftlicher Vereins in Niederlandisch Indien in Batavia.

Die Echinodermen. 1. Holothuroidea. Natuurkundig Tijdschrift

Nederlandische Indie 47: 181-220,2 pis.

Sluiter, C. P. 1901. Siboga -Expedite. Die Holothurien der Siboga-

Expedition 44. 142 pp, 11 pis.

Smirnov, A. V. 1989. A new species of holothurians Trochodota

inexspectata (Synaptida, Chiridotidae) from the Simushir Island

(Kuril Islands). Zoologicheskii Zhurnal 68(6): 156-160.

Smirnov, A. V. 2012. System of the class Holothuroidea. Paleontological

Journal 46(8): 793-832.

Stimpson, W. 1855. Descriptions of some new marine invertebrata.

Proceedings of the Academy of Natural Sciences, Philadelphia 7(10):

385-395

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Tamura, K., Stecher, G., Peterson, D., Filipski, A., & Kumar, S. 2013.

MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0.

Molecular Biology and Evolution 30: 2725-2729.

Thandar, A. S. 1989. The sclerodactylid holothurians of southern

Africa, with the erection of one new subfamily and two new

genera. South African Journal Zoology 24(4): 290-304.

Theel, H. 1886. Report on the Holothurioidea dredged by H.M.S.

Challenger during the years 1873-1876. Report on the scientific

results of the voyage of H.M.S. Challenger, Zoology 14 (39):

1-290, 16 pis.

Troschel, F. H. 1846. Neue Holothurien-Gattungen. Archiv fiir

Naturgeschichte 12(1): 60-66.

Appendix 1. Tissue samples for genetic data from Camden Sound sea cucumbers.

Registration

Barcode

Type

Tissue

Taxa

WAM

CSIRO

MOLAF

Z89000

10000043

paratype

1759

Holothuria (Metriatyla) keesingi O’Loughlin sp. nov.

Z89001

10000403

paratype

1760

Holothuria (Metriatyla) keesingi O’Loughlin sp. nov.

Z89002

10001168

paratype

1761

Holothuria (Metriatyla) keesingi O’Loughlin sp. nov.

Z89003

10001320

paratype

1762

Holothuria (Metriatyla) keesingi O’Loughlin sp. nov.

Z89004

10001821

paratype

1763

Holothuria (Metriatyla) keesingi O’Loughlin sp. nov.

Z89005

10001954

paratype

1764

Holothuria (Metriatyla) keesingi O’Loughlin sp. nov.

Z89006

10002938

holotype

1765

Holothuria (Metriatyla) keesingi O’Loughlin sp. nov.

Z89007

10002963

paratype

1766

Holothuria (Metriatyla) keesingi O’Loughlin sp. nov.

Z89008

No no.

1767

Holothuria (Thymiosycia) gracilis (Semper, 1868)

Z89009

10003037

1768

Stichopus unresolved species complex including Stichopus herrmanni Semper,

1868

Z89010

10003038

1769

Stichopus unresolved species complex including Stichopus herrmanni Semper,

1868

Z89011

10001261

1770

Globosita elnazae O’Loughlin, 2014 (in O’Loughlin, Mackenzie &

VandenSpiegel, 2014)

Z89012

10001262

1771

Globosita elnazae O’Loughlin, 2014 (in O’Loughlin, Mackenzie &

VandenSpiegel, 2014)

Z89013

10002087

1772

Cercodemas anceps Selenka, 1867

Z89014

10000001

1773

Colochirus quadrangularis Troschel, 1846

Z89015

10000582

1692

Colochirus quadrangularis Troschel, 1846

Z89016

10000647

1774

Colochirus quadrangularis Troschel, 1846

Z89017

10001114

1775

Colochirus quadrangularis Troschel, 1846

Z89018

10001331

1776

Colochirus quadrangularis Troschel, 1846

Z89019

10001420

1777

Colochirus quadrangularis Troschel, 1846

Z89020

10001560

1778

Colochirus quadrangularis Troschel, 1846

Z89021

10001767

1690

Colochirus quadrangularis Troschel, 1846

Z89022

10001971

1691

Colochirus quadrangularis Troschel, 1846

Z89023

10002500

1779

Colochirus quadrangularis Troschel, 1846

Z89024

10002692

1780

Colochirus quadrangularis Troschel, 1846

Z89025

10002871

1781

Colochirus quadrangularis Troschel, 1846

Z89026

10002268

holotype

1688

Plesiocolochirus minaeus O’Loughlin sp. nov.

Z89027

53713

1782

Leptopentacta grisea H. L. Clark, 1938

Z89028

10001662

1783

Plesiocolochirus sp. 1, unresolved species complex including P. australis

(Ludwig, 1875)

Z89029

10001822

1784

Plesiocolochirus sp. 1, unresolved species complex including P. australis

(Ludwig, 1875)

Z89030

10002101

1785

Plesiocolochirus sp. 1, unresolved species complex including P. australis

(Ludwig, 1875)

Z89031

10000760

1786

Pseudocolochirus axiologus (H. L. Clark, 1914)

Z89032

10002389

1787

Pseudocolochirus axiologus (H. L. Clark, 1914)

Z89033

10001306

1788

Phyllophorus (Phyllophorella) spiculata Chang, 1935

P.M. O’Loughlin, C. Harding & G. Paulay

Registration

Barcode

Type

Tissue

Taxa

WAM

CSIRO

MOLAF

Z89034

10000113

1789

Phyllophorus (Urodemella) holothurioides Ludwig, 1875

Z89035

10001277

1790

Phyllophorus (Urodemella) holothurioides Ludwig, 1875

Z89036

10000277

1791

Havelockia versicolor (Semper, 1867)

Z89037

10000776

1792

Havelockia versicolor (Semper, 1867)

Z89038

10001220

1793

Havelockia versicolor (Semper, 1867)

Z89039

10001124

1794

Havelockia versicolor (Semper, 1867)

Z89040

10001321

1795

Havelockia versicolor (Semper, 1867)

Z89041

10001417

1796

Havelockia versicolor (Semper, 1867)

Z89042

10001637

1797

Havelockia versicolor (Semper, 1867)

Z89043

10001924

1798

Havelockia versicolor (Semper, 1867)

Z89044

10002314

1799

Havelockia versicolor (Semper, 1867)

Z89045

10002323

1800

Havelockia versicolor (Semper, 1867)

Z89046

10002688

1801

Havelockia versicolor (Semper, 1867)

Z89047

10002689

1802

Havelockia versicolor (Semper, 1867)

Z89048

10002752

1803

Havelockia versicolor (Semper, 1867)

Z89049

10002838

1804

Havelockia versicolor (Semper, 1867)

Z89050

10002939

1805

Havelockia versicolor (Semper, 1867)

Z89051

10002243

1806

Massinium bonapartum O’Loughlin, 2014 (in O’Loughlin, Mackenzie &

VandenSpiegel, 2014)

Z89052

10002725

1807

Massinium bonapartum O’Loughlin, 2014 (in O’Loughlin, Mackenzie &

VandenSpiegel, 2014)

Z89053

10002966

1808

Massinium bonapartum O’Loughlin, 2014 (in O’Loughlin, Mackenzie &

VandenSpiegel, 2014)

Z89054

10002958

holotype

1809

Neothyonidiumip .) insolitum O’Loughlin sp. nov.

Z89055

10001531

1810

Hemithyone semperi (Bell, 1884)

Z89056

10002195

1811

Stolus canescens (Semper, 1867)

Z89057

10001097

1812

Thy one papuensis Theel, 1886

Z89058

10001259

1813

Thy one papuensis Theel, 1886

Z89059

10002102

1814

Thy one papuensis Theel, 1886

Z89060

10002542

1815

Thy one papuensis Theel, 1886

Z89061

10000683

1816

Actinocucumis longipedes H. L. Clark, 1938

Z89062

10000111

1817

Actinocucumis solanderi O’Loughlin, 2014 (in O’Loughlin, Mackenzie &

VandenSpiegel, 2014)

Z89063

10001112

1818

Actinocucumis solanderi O’Loughlin, 2014 (in O’Loughlin, Mackenzie &

VandenSpiegel, 2014)

Z89064

10002832

1819

Actinocucumis solanderi O’Loughlin, 2014 (in O’Loughlin, Mackenzie &

VandenSpiegel, 2014)

Z89065

10002613

1820

Actinocucumis typica Ludwig, 1875

Z89066

10002792

1821

Actinocucumis typica Ludwig, 1875

Z89067

10000685

1822

Mensamaria intercedens (Lampert, 1885)

Z89068

10001307

1823

Mensamaria intercedens (Lampert, 1885)

Z89069

53720

1824

Molpadia scabrum (Sluiter, 1901)

Z89070

10000038

1825

Protankyra insolens (Theel, 1886)

Z89071

10000142

holotype

1826

Protankyra torquea O’Loughlin sp. nov.

Z89072

10002967

1827

Protankyra verrilli (Theel, 1886)

Z89073

10001534

1828

Synaptula lamperti Heding, 1928

Z89074

10000777

1829

Synaptula recta (Semper, 1867)

Z89075

10000919

1830

Synaptula recta (Semper, 1867)

Z89076

10002614

1831

indeterminate unknown species (ring and tentacles eviscerated)

The sea cucumbers of Camden Sound in northwest Australia, including four new species (Echinodermata: Holothuroidea)

Appendix 2. List of tissues (with tissue sample code numbers, specimen repositories, specimen registration numbers, specimen source locations,

and GenBank Accession numbers) from Colochirus Troschel, 1846 and Plesiocolochirus Cherbonnier, 1946 specimens with sequences in the

phylogenetic tree.

Genus

species

Tissue code

Museum

Registration

Location

GenBank

Colochirus

species 1 GP

UF 10059

10059

Heron Island

KX844560

Colochirus

species 1 GP

MOLAF 1263

G22502 (1)

Lizard Island

KX844598

Colochirus

species 1 GP

MOLAF 1264

G22502 (2)

Lizard Island

KX844574

Colochirus

species 1 GP

MOLAF 1265

G22502 (3)

Lizard Island

KX844583

Colochirus

species 1 GP

UF 8504

UF8504

Lizard Island

KX844613

Colochirus

species 1 GP

UF 2249

UF2249

PNG

KX844564

Colochirus

species 1 GP

UF 10918

UF10918

Okinawa

KX844566

Colochirus

species 1 GP

PH-28

17852

Philippines

KX844572

Colochirus

species 1 GP

PH-34

17853

Philippines

KX844573

Colochirus

species 1 GP

UF 957

957

Okinawa

KX844569

Colochirus

species 1 GP

MOLAF 1200

WAM

Z26234 (11)

Kimberley

KX844568

Colochirus

species 1 GP

UF 7400

7400

Madagascar

KX844611

Colochirus

species 1 GP

UF 7510

7510

Madagascar

KX844609

Colochirus

quadrangularis

MOLAF 1693

NMV

F201782

N Australia

KX844576

Colochirus

quadrangularis

UF 13683

13683

Singapore

KX844595

Colochirus

quadrangularis

UF 13667

13667

Singapore

KX844565

Colochirus

quadrangularis

MOLAF 1692

WAM

Z89015

Kimberley

KX844610

Colochirus

quadrangularis

MOLAF 398

NMV

F149742

Kimberley

KX844559

Colochirus

quadrangularis

MOLAF 1691

WAM

Z89022

Kimberley

KX844596

Colochirus

quadrangularis

MOLAF 1690

WAM

Z89021

Kimberley

KX844594

Colochirus

quadrangularis

MOLAF 1453

WAM

Z27858

Kimberley

KX844605

Colochirus

quadrangularis

MOLAF 1209

NMV

F173259

N Australia

KX844601

Colochirus

quadrangularis

MOLAF 1210

NMV

F173260

N Australia

KX844562

Colochirus

quadrangularis

QM09 058

SBD503854

Queensland

KX844586

Colochirus

quadrangularis

QM09 074

TS80000215

Queensland

KX844577

Colochirus

robustus

MOLAF 396

NMV

F149737

Kimberley

KX844579

Colochirus

robustus

MOLAF 397

NMV

F149738

Kimberley

KX844606

Colochirus

robustus

UF 17373

17373

Philippines

KX844561

Colochirus

robustus

UF 17672

17672

Philippines

KX844582

Plesiocolochirus

challengeri

MOLAF 1460

NMV

F203000

Kimberley

KX844597

Plesiocolochirus

challengeri

MOLAF 1461

WAM

Z27862

Kimberley

KX844570

Plesiocolochirus

challengeri

MOLAF 1213

NMV

F173263

N Australia

KX844587

Plesiocolochirus

species 2 GP

UF 10077

10077

Heron Island

KX844571

Plesiocolochirus

species 2 GP

UF 9986

9986

Heron Island

KX844563

Plesiocolochirus

species 2 GP

UF 10041

10041

Heron Island

KX844607

Plesiocolochirus

tessellarus

MRAC 2005 39

MRAC

2616

Comoros

KX844590

Plesiocolochirus

species 1 GP

MOLAF 394

NMV

FI 50795

Kimberley

KX844567

Plesiocolochirus

species 1 GP

MOLAF 395

NMV

FI 50805

Kimberley

KX844604

Plesiocolochirus

species 1 GP

MOLAF 1444

WAM

Z27854 (1)

Kimberley

KX844584

Plesiocolochirus

species 1 GP

MOLAF 1443

WAM

Z27854 (2)

Kimberley

KX844578

Plesiocolochirus

species 1 GP

MOLAF 1196

WAM

Z26229 (3)

Kimberley

KX844585

Plesiocolochirus

species 1 GP

UF 8952A

8952A

Palau

KX844589

Plesiocolochirus

species 1 GP

UF 9535

9535

Ningaloo

KX844612

Plesiocolochirus

species 1 GP

UF 17730

17730

Japan

KX844575

Plesiocolochirus

minaeus sp. nov.

MOLAF 1688

WAM

Z89026

Kimberley

KX844602

P.M. O’Loughlin, C. Harding & G. Paulay

Genus

species

Tissue code

Museum

Registration

Location

GenBank

Plesiocolochirus

ignavus

MOL AF 430

NMV

FI 51840 (1)

Victoria

KX844592

Plesiocolochirus

ignavus

MOL AF 431

NMV

FI 51840 (2)

Victoria

KX844603

Plesiocolochirus

ignavus

MOLAF 1175

NMV

F173252

Victoria

KX844593

Plesiocolochirus

ignavus

MOLAF 1177

NMV

F173255 (2)

Victoria

KX844588

Plesiocolochirus

ignavus

MOLAF 453

NMV

FI25377

Victoria

KX844591

Plesiocolochirus

ignavus

MOLAF 454

NMV

FI 25376

Victoria

KX844580

Plesiocolochirus

ignavus

MOLAF 450

NMV

FI 26892 (1)

Victoria

KX844600

Plesiocolochirus

ignavus

MOLAF 451

NMV

FI 26892 (2)

Victoria

KX844581

Plesiocolochirus

ignavus

MOLAF 452

NMV

FI 26892 (3)

Victoria

KX844599

Memoirs of Museum Victoria 75:53-70 (2016) Published 2016

1447-2554 (On-line)

http://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

The Discovery Expedition sea cucumbers (Echinodermata: Holothuroidea)

(http://zoobank.org/urn:lsid:zoobank.org:pub:7039F593-8FE2-4668-BBC9-E18F2D82F8F8)

P. MARK O’LoUGHLIN 1 * (http://zoobank.org/urn:lsid:zoobank.org:author:97B95F20-36CE-4A76-9DlB-26A59FBCCE88),

ELNAZ TAVANCHEH 1 (http://zoobank.Org/urn:lsid:zoobank.org:author:152088AC-CBD6-4913-931C-7180818DD345) AND

CAROLINE Harding 1 (http://zoobank.org/urn:lsid:zoobank.org:author:FC3B4738-4973-4A74-B6A4-F0E606627674)

1 Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, Victoria 3001, Australia

* To whom correspondence should be addressed. E-mail: pmoloughlin@edmundrice.org

Abstract O’Loughlin P. M., Tavancheh, E. & Harding, C. 2016. The Discovery Expedition sea cucumbers (Echinodermata:

Holothuroidea). Memoirs of Museum Victoria 75: 53-70.

Identifications of all lots of Holothuroidea specimens collected from February 1926 to January 1939 by the

Discovery Expedition are listed with station data, locality and depth. This report includes identifications reported

previously by Heding (in Heding & Panning), O’Loughlin & Ahearn, O’Loughlin et al. and O’Loughlin & VandenSpiegel.

New taxa from the Discovery Expedition have been reported previously, and a summary is provided. The new taxa herein

are for equatorial West Africa specimens: new genus Cucusquama O’Loughlin and new species Cucusquama wesafrica

O’Loughlin. Systematic notes are provided for genera Clarkiella Heding (in Heding & Panning) and Echinopsolus Gutt,

and species Ocnus capensis (Theel), genera Parathyonidium Heding (in Heding & Panning) and Pentactella Verrill, and

species Psolus dubiosus Ludwig & Heding, Psolus lockhartae O’Loughlin & Whitfield and Sigmodota contorta (Ludwig).

Earlier echinoderm specialists who studied Discovery Expedition holothuroids are acknowledged: Cynthia Ahearn,

Elizabeth Deichmann, Svend Heding, Melanie Mackenzie, Albert Panning and Emily Whitfield.

Keywords Falkland Islands, New Zealand, Ross Sea, South Georgia, West Africa, Clarkiella, Echinopsolus, Parathyonidium,

Pentactella, Psolus, new genus, new species.

Introduction

In 1918 an Interdepartmental Committee on Research and

Development in the Falkland Islands Dependencies was

formed. Its report to the British Parliament in 1920 resulted in

the appointment of a Discovery Committee in 1923. The first

Discovery Expedition sailed south on 24 September 1925

aboard Robert Falcon Scott’s Discovery as its research vessel

(Figure la). It had been built for Scott’s 1901-1904 British

National Antarctic Expedition. In the Falklands it was found

to be unsuitable for scientific operations in open waters. The

William Scoresby (Figure 2) was built in 1925 and was used

principally in whale research. A marine biological station

(known as Discovery House ) was built at King Edward Point

on South Georgia. In 1927 the Discovery II (Figure lb) was

built and was judged to be an outstanding success. The

Discovery Committee was dissolved in 1949 when it became

part of the National Institute of Oceanography, located in

Surrey, UK. The Whale Research Unit moved, briefly, to the

Natural History Museum in London. It later became part of

the British Antarctic Survey (BAS), based in Cambridge. In

1995 the Institute of Oceanographic Sciences became part of

the Southampton Oceanography Centre. Scott’s second

Antarctic expedition on the Terra Nova (1910-1913) collected

marine invertebrate animals amongst which was a large

collection of sea cucumbers. This Terra Nova sea cucumber

collection is referred to below.

The results of the Discovery Expedition have been

documented in the 37 volumes of Discovery Reports (1928-

1980). The ships used in the early decades of the Expedition

were the RRS Discovery I (Figure la; from 25 September

1925 to 1927), RRS Discovery II (Figure lb; five voyages

from 1929 to 1935, and sixth voyage in 1950), and RRS

William Scoresby (Figure 2, eight voyages from 1926 to

1951). The localities visited by the Expedition were

predominantly the Falkland Islands and Falkland Islands

Dependencies (South Georgia, S. Sandwich Is, S. Shetland Is,

S. Orkney Is) and the Ross Sea. Some collections were made

from the Marine Biological Station {Discovery House ) at

King Edward Point, South Georgia. Some collections were

made during voyages to and from the Falkland Islands. Thus

species identifications are included here for Antipodes I.,

Balleny I., the Haakon VII Sea, Marion I., New Zealand, the

Ross Sea, Marine Station 82 in Saldanha Bay in south-west

Africa, and equatorial west Africa.

P.M. O’Loughlin, E. Tavancheh & C. Harding

Figure 1. Ships from which the early Discovery Expedition was conducted, a, Scott’s RRS Discovery I (collected from 25 Sep 1925 to 1927); b,

RRS Discovery II (collected for five voyages from 1929 to 1935, and a sixth voyage in 1950).

The Discovery Expedition to the South Sandwich Islands in

1927 was not conducted by RRS Discovery or RRS William

Scoresby, and is reported in Discovery Report Vol. 3. The

Discovery Expedition to the Ross Sea from Nov. 1928 to Feb.

1929 was not conducted by RRS Discovery or RRS William

Scoresby , and is reported in Discovery Report Vol. 3. There are

no holothuroid lots from these expeditions in the collection

reported here. The British Antarctic Survey commenced its

survey of the Falkland Islands Dependencies in 1943, and the

BAS surveys continue to the present day.

Three papers have been published in the Discovery Reports

on the early echinoderm collections. Mortensen (1936, Vol. 12)

reported on the Discovery Echinoidea and Ophiuroidea

collected from 1925 to 1935. Dilwyn John (1938, in Vol. 18)

reported on the Discovery Crinoidea collected from 1935 to

1937. Fisher (1940, in Vol. 20 issued in 1941) reported on the

Discovery Asteroidea collected from 1925 to 1936. Some

Discovery Expedition Holothuroidea have been reported in

Heding & Panning (1954), O’Loughlin & Ahearn (2008),

O’Loughlin & VandenSpiegel (2010) and O’Loughlin et al.

(2014). This paper includes these taxa reported previously, and

is the first comprehensive Discovery report on the Holothuroidea

that were collected from 28 February 1926 to 30 January 1939.

The Danish holothuroid specialist Svend Geisler Heding

(1902-1949, see Acknowledgments, Figure 3a) studied the

Discovery Expedition holothurians in the Zoological Museum in

the University of Copenhagen (ZMUC) until his death. He

described two new genera and species: Clarkiella discoveryi

Heding (in Heding & Panning 1954); Parathyonidium incertum

Heding (in Heding & Panning 1954). These new taxa were

ascribed to Heding posthumously by the German sea cucumber

specialist Albert Panning (1894-1978, Figure 3b). Panning

included the descriptions in Heding & Panning 1954 from

Heding’s notes and mentioned that he did this in collaboration

with Elizabeth Deichmann (working at that time on the Discovery

Expedition holothuroids in the Museum of Comparative Zoology

at Harvard). The Heding types are lodged in London (NHMUK),

Paris (MNHN) and Copenhagen (ZMUC) (see Table 2 below).

The Discovery Expedition sea cucumbers (Echinodermata: Holothuroidea)

Figure 2. RRS William Scoresby served the Discovery Expedition for eight voyages from 1926 to 1951.

Shortly after Heding’s death in 1949, H. W. Parker (the

Keeper of Zoology in the British Museum) wrote on 25 th July

in 1950 to Elizabeth Deichmann (c/o F. Jensenius Madsen in

the Zoology Museum in the University of Copenhagen)

inviting her to work on the holothurians in the Terra Nova ,

BANZARE and John Murray collections. The letter noted that

these collections belonged to the British Museum. She was

also invited to consider contributing to the “Reports of the

John Murray Expedition”. The letter of invitation anticipated

that Dr. Macintosh (in the National Institute of Oceanography

that managed the Discovery collections) would probably also

like Deichmann to study the Discovery Expedition

holothurians and contribute to the Discovery Reports. These

four collections were still in Copenhagen at the time. The

letter notes that Dr. Madsen had the notes left by Heding.

These notes were evidently passed on to Deichmann and were

presumably the source from which Albert Panning published

the new taxa that he ascribed to Heding. Shortly after this

communication the collections were returned to the British

Museum in London.

Elizabeth Deichmann (1896-1975, Figure 4a), like Heding,

was Danish and a protegee of echinoderm specialist Theodor

Mortensen (1868-1952). In 1929 Deichmann began working

in The Museum of Comparative Zoology in Harvard University

in Massachusetts. In response to the invitation to work on the

four major holothuroid collections she visited London in 1950

to study the collections. A National Institute of Oceanography

internal letter (David Pawson pers. comm.) of 7 August 1963,

directed to Dr Macintosh, noted: “In Miss Ailsa Clark’s

absence in America (Miss Clark is in charge of Echinoderms),

Dr Deichmann arranged with an assistant in the department to

have a considerable part of the material packed up and sent

over to Harvard. Unfortunately no list was made of the station

numbers and species selected by Dr Deichmann, but I have

asked Miss Clark to check what remains of the Discovery

specimens in her department against my list of material

originally sent to Heding. I enclose the full data on a separate

sheet”. This letter noted that the last communication with

Deichmann was in 1961, and at the time of this communication

(1963) she had retired from work at the MCZ, Harvard

P.M. O’Loughlin, E. Tavancheh & C. Harding

Figure 3. Authors who published the first new sea cucumber genera and species from the Discovery Expedition in their monograph on the

Phyllophoridae in 1954. a, Danish holothuroid specialist Svend Geisler Heding (1902-1949; photo provided by Tom Schioette in ZMUC); b,

German holothuroid specialist Albert Panning (1894-1978; photo provided by David Pawson in USNM).

University. But in a letter to David Pawson ( pers. comm) on

25 February 1967 it appears that she was still working at the

MCZ and hoped to work with Pawson on the Discovery

holothuroids. This did not eventuate. Deichmann published

her last paper in 1958. None of her papers mentions any

Discovery Expedition holothuroids.

The collections arrived at the MCZ in 1950. In a letter

dated 25 th February 1967 to David Pawson (Smithsonian

Institution, pers. comm), Deichmann commented that the

collection was in poor condition when she received it and that

localities were “somewhat mixed up”. But she wrote that she

“got good help from Ailsa Clark” to resolve issues. Elizabeth

further commented that she had made good headway and had

a lot of notes about the genus Psolus. There was some evidence

of Deichmann’s work when we received the collection in

Museum Victoria (2008). Deichmann did not publish on any

of the taxa or contribute to any report. Unfortunately we do

not have her notes on Psolus.

Following Deichmann’s retirement the Discovery and

Terra Nova collections were transported by David Pawson

{pers. comm, probably in 1973) from the MCZ to the United

States National Museum in Washington (Smithsonian

Institution). In the Smithsonian an echinoderm specialist

Cynthia (Gust) Ahearn (1952-2008, Figure 4b) identified many

lots personally, and then collaboratively with Mark O’Loughlin.

O’Loughlin & Ahearn (2008) included numerous lots of

Discovery Expedition psolid species (see Table 3 below).

Ahearn had the Discovery holothuroid collection sent to

Museum Victoria in 2008, with permission from the Natural

History Museum (UK). Mark O’Loughlin, Melanie Mackenie

(Figure 4c) and Emily Whitfield (Figure 4d) then continued

with identifications. This work was delayed in 2010 until

related systematic issues were resolved. But O’Loughlin &

VandenSpiegel (2010) included numerous Discovery synaptids

(as apodids) in their comprehensive review (see Table 4 below).

Identification by O’Loughlin in Museum Victoria

recommenced in 2015, and identifications of the Discovery

holothuroid lots have now been completed and earlier

identifications changed or confirmed. The results are collated

and reported here (see Table 1 below). Apart from the Heding

types lodged in Copenhagen (ZMUC) and Paris (MNHN), the

Discovery holothuroids are lodged in the NHMUK.

The Discovery Expedition holothuroid lots that are the

subject of this report were collected in the 1920’s and 1930’s.

The collection has been transported on loan numbers of times,

and studied by several specialists. Today, the collection is in only

fair condition in terms of quality of preservation and clarity of

labelling. Apart from the thorough work of Cynthia Ahearn in

the Smithsonian there were very few attempted identification

labels in the lots. David Pawson {pers. comm) commented that

he noticed that very few lots had labels when he transported the

collection from Harvard to Washington. We found that the

condition of some specimens was such that we could not identify

them. We note again the National Institute of Oceanography

The Discovery Expedition sea cucumbers (Echinodermata: Holothuroidea)

Figure 4. Sea cucumber specialists who contributed to the identifications of the specimen lots that are the subject of this report, a, Elizabeth

Deichmann (1896-1975; photo provided by David Pawson in USNM); b, Cynthia (Gust) Ahearn (1952-2008); c, Melanie Mackenzie (Museum

Victoria); d, Emily Whitfield (volunteer in Museum Victoria).

P.M. O’Loughlin, E. Tavancheh & C. Harding

reported that Deichmann had “a considerable part of the material

packed up and sent over to Harvard”, and Deichmann’s comment

that some of the locality data were “somewhat mixed up”. We

found some discrepancies between the data on labels in lots and

the lists of data in the Discovery Reports. We thus have some

reservations about the absolute accuracy of what we are reporting

here.

The Terra Nova (1910-1913) collection was sent to Museum

Victoria from the Smithsonian with the Discovery collection,

and the lots have been identified where possible. This collection

and the documentation are in poor condition. The collection

currently in Museum Victoria is being prepared for return to

the Natural History Museum in London. We are collaborating

with Stefano Schiaparelli ( pers. comm. University of Genova)

who is developing a report on the holothuroids of Terra Nova

Bay that will incorporate some of the Terra Nova collection

data. O’Loughlin (2009) published a final report on The British,

Australian and New Zealand Antarctic Research Expedition

holothuroids (BANZARE, 1929-1930). This collection is

lodged in the South Australian Museum.

body wall, while the second subfamily Colochirinae Panning,

1949 has plates and bowl/cup/basket ossicles in the body wall.

Cucusquama O’Loughlin gen. nov.

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act:

0BE9B60F-195B-4F04-A06A-7EAF0BAA9D44

Diagnosis. Cucumariinid genus; body sub-pentagonal in

transverse section, fusiform; tentacles 10, short, lobed;

calcareous ring cucumariid-like; complete calcareous body

cover of imbricating, single-layered, perforated plates/scales,

free ends point posteriorly; tube feet radial only, walls covered

with scales; absence of cups and tables.

Type species and locality. Cucusquama wesafrica O’Loughlin

sp. nov. (equatorial west Africa). Monotypic.

Etymology. Named Cucu from the family name Cucumariidae,

with recognition of cucumariid like characters, and squama

from the Latin squama (meaning scale) with reference to the

body cover of imbricating scales.

Abbreviations

BAS British Antarctic Survey

D Station data prefix for RRS Discovery 1 and II

collections

MCZ Museum of Comparative Zoology, Harvard

University

MS Station data prefix for the Marine Biological

Stations in South Georgia and in South Africa

MNHN Museum national d’Histoire naturelle, Paris

NHMUK Natural History Museum, London

NMV Museum Victoria, Australia

USNM United States National Museum (part of

Smithsonian Institution)

WS Station data prefix for RRS William Scoresby

collections

ZMUC Zoological Museum University of Copenhagen

Methods

The macro images of preserved specimens were taken by

Caroline Harding, with Mark O’Loughlin, using a Canon 5D

mark ii camera mounted on a camlift Visionary Digital auto

stepper, and Zerene Image Stacker, Adobe Lightroom and

Photoshop for image processing and editing. The photos of

ossicles were taken by Caroline Harding, with Mark

O’Loughlin, using a LEICA DM5000 B microscope, Leica

application software, and Helicon Focus montage software.

New taxa from the Discovery Expedition (1926-1939)

Order Dendrochirotida Grube, 1840

Family Cucumariidae Ludwig, 1894

Subfamily Cucumariinae Ludwig, 1894 sensu Panning 1949

Remarks. The subfamily Cucumariinae has plates only in the

Remarks. The sub-pentagonal form, complete body cover of

imbricating scales, 10 short lobed tentacles, and radial series of

tube feet is a unique combination of morphological characters

within family Cucumariidae. We reluctantly establish a new

monotypic genus.

Cucusquama wesafrica O’Loughlin sp. nov.

Zoobank LSID. http://z 00 bank. 0 rg/urn:lsid:z 00 bank. 0 rg:act:

D4A442DB-36E7-453C-945D-47FA23AB6CAA

Table 1; figures 5, 6

Material examined. Holotype. West Africa, Luanda, Angola,

Discovery stn D 274, -8.84 13.23 64-65 m, 4 Aug 1927, NHMUK

2016.148.

Paratypes (4). West Africa, off Cape Lopez, Gambon (French

Congo), Discovery stn D 279, -0.63 8.70 58-67 m, 10 Aug 1927,

NHMUK 2016.149-152.

Description. Body (preserved) up to 13 mm axial length, up to

3 mm high, body form sub-pentagonal, body tapered anteriorly

and posteriorly, long posterior taper to create a tail; body

completely invested in imbricating scales, free ends of

imbricating scales point posteriorly; oral disc at base of oral

cavity created by anteriorly-projecting scales; no anal scales

detected; 10 short, lobed, black tentacles in ring on oral disc;

ring not strongly calcified, form weakly evident, cucumariid-

like; internal organs shriveled, brittle; tube feet on radii only, in

single spaced series, up to 12 tube feet on any radius, tube feet

more strongly developed on mid-ventral and ventro-lateral

radii, wall of tube feet covered with imbricating scales.

Body wall ossicles single-layered perforated plates only,

irregularly oval; large plates scales up to 600 pm long; some

irregular smaller plates up to about 170 pm long; no evidence

of cups or tables detected. No tube foot endplates detected. No

tentacle ossicles detected.

Distribution. Equatorial West Africa, off Angola and Gambon,

58-67 m.

The Discovery Expedition sea cucumbers (Echinodermata: Holothuroidea)

Table 1. Complete list of Discovery Expedition taxa collected from 1926 to 1939 (H indicates holotype; P indicates paratype(-s)).

Taxa (grouped in orders, in alphabetical sequence)

Station

number

Locality

Depth

NHMUK registration

(+ MNHN, ZMUC)

Aspidochirotida

Pseudostichopus spiculiferus (O’Loughlin, 2002)

D 1658

Ross Sea

520 m

2011.169-170 (3)

Pseudostichopus spiculiferus (O’Loughlin, 2002)

D 600

South Shetland Islands

501-527 m

2011.297(1)

Pseudostichopus spiculiferus (O’Loughlin, 2002)

D 1660

Ross Sea

351 m

2016.175-177

Dendrochirotida

Amphicyclus thornsoni (Hutton, 1878)

D 941

Cook Strait, New Zealand

128 m

2016.134-135

Cladodactyla crocea (Lesson, 1830)

WS 869

Falkland Islands

187 m

2016.100

Cladodactyla crocea (Lesson, 1830)

WS 231

Falkland Islands

159-167 m

2016.101

Cladodactyla crocea (Lesson, 1830)

WS 867

Falkland Islands

147-150 m

2016.102

Cladodactyla crocea (Lesson, 1830)

WS 804

Falkland Islands

143-150 m

2016.147

'Clarkiella discoveryi Heding, 1954 (in Heding &

Panning, 1954) H

D 474

W of Shag Rocks, South

Georgia

199 m

ZMUC HOL-64

l Clarkiella discoveryi Heding, 1954 (in Heding &

Panning, 1954) P

D 474

W of Shag Rocks, South

Georgia

199 m

ZMUC HOL-247

Crucella scotiae (Vaney, 1906)

D 2567

Haakon VII Sea

100-124 m

2011.196

Crucella scotiae (Vaney, 1906)

D 175

South Shetland Islands

200 m

2011.168

Cucamba psolidiformis (Vaney, 1908)

D 1872

South Shetland Islands

247 m

2016.140

Cucamba psolidiformis (Vaney, 1908)

D 1660

Ross Sea

351 m

2016.145

Cucumaria dudexa O’Loughlin & Manjon-Cabeza,

2009b

D 456

Ross Sea

40-45 m

2011.141-150(16)

Cucusquama wesafrica O’Loughlin sp. nov. H

D 274

off Luanda, Angola

64—65 m

2016.148

Cucusquama wesafrica O’Loughlin sp. nov. P

D 279

Cape Lopez, French Congo

58-67 m

2016.149-152

Echinopsolus acanthocola Gutt, 1990

D 1660

Ross Sea

351 m

2016.141

Echinopsolus acanthocola Gutt, 1990

D 1660

Ross Sea

351 m

2016.156-161

Echinopsolus acutus (Massin, 1992)

D 1652

Ross Sea

567 m

2011.341

Echinopsolus acutus (Massin, 1992)

D 1872

South Shetland Islands

247 m

2011.342-344

Echinopsolus acutus (Massin, 1992)

D 1652

Ross Sea

567 m

2016.162-170

Echinopsolus attenuatus (Vaney, 1906)

D 1652

Ross Sea

567 m

2011.366

Echinopsolus georgiana (Lampert, 1886) group

Gutt, 1988

D 1652

Ross Sea

567 m

2011.367

Echinopsolus georgiana (Lampert, 1886) group

Gutt, 1988

WS 228

Falkland Islands

229-236 m

2016.154

Echinopsolus koehleri (Vaney, 1914)

WS 245

Falkland Islands

209-304 m

2016.171-173

Echinopsolus koehleri (Vaney, 1914)

D 159

South Georgia

160 m

2011.314-315

Echinopsolus koehleri (Vaney, 1914)

D 160

Shag Rocks

0-180 m

2011.316-320

Echinopsolus koehleri (Vaney, 1914)

WS 33

South Georgia

0-130 m

2011.321

Echinopsolus koehleri (Vaney, 1914)

WS 33

South Georgia

0-130 m

2011.322

Echinopsolus koehleri (Vaney, 1914)

D 156

South Georgia

200-236 m

2011.323

Echinopsolus koehleri (Vaney, 1914)

D 148

South Georgia

132-148 m

2011.324

Echinopsolus koehleri (Vaney, 1914)

WS 840

Falkland Islands

368-463 m

2011.325

Echinopsolus koehleri (Vaney, 1914)

D 175

South Shetland Islands

200 m

2011.326

Echinopsolus koehleri (Vaney, 1914)

D 363

South Sandwich Islands

278-329 m

2011.327-328

Echinopsolus koehleri (Vaney, 1914)

D 160

Shag Rocks

0-180 m

2011.329-336

Echinopsolus mollis (Ludwig & Heding, 1935)

D 363

South Sandwich Islands

278-325 m

2016.104—107

Hemioedema spectabilis (Ludwig, 1883)

WS 797

Falkland Islands

112-114 m

2016.136

Heterocucumis steineni (Ludwig, 1898)

D 456

Ross Sea

40-45 m

2011.162-167

Heterocucumis steineni (Ludwig, 1898)

D 2567

Haakon VII Sea

100-124 m

2011.197

P.M. O’Loughlin, E. Tavancheh & C. Harding

Taxa (grouped in orders, in alphabetical sequence)

Station

number

Locality

Depth

NHMUK registration

(+ MNHN, ZMUC)

Heterothyone ocnoides (Dendy, 1897)

D 939

off Dargaville, New

Zealand

87 m

2016.113-115

Neopsolidium convergens (Herouard, 1901)

D 724

Strait of Magellan

0-5 m

2011.126-132

Neopsolidium convergens (Herouard, 1901)

WS 84

Falkland Islands

74—75 m

2011.133

Neopsolidium convergens (Herouard, 1901)

D 51

Falkland Islands

115 m

2011.134-137

Neopsolidium kerguelensis (Theel, 1886)

D 1564

Marion Island

108-113 m

2016.178

Neothyonidium armatum Pawson, 1965

D 941

Cook Strait, New Zealand

128 m

2016.153

Ocnus capensis (Theel, 1886)

MS 82

Saldanha Bay, South Africa

7-14 m

2016.143

Paracucumis turricata (Vaney, 1906)

D 1651

Ross Sea

594 m

2011.139-140

2 Parathyonidium incertum Heding, 1954 (in

Heding & Panning, 1954) H

2 D 170

2 South Shetland Islands

2 342 m

3 ZMUC HOL-93

Parathyonidium incertum Heding, 1954 (in

Heding & Panning, 1954) P

D 170

South Shetland Islands

342 m

ZMUC HOL-300

Parathyonidium incertum Heding, 1954 (in

Heding & Panning, 1954) P

D 170

South Shetland Islands

342 m

2011.171-173

Parathyonidium incertum Heding, 1954 (in

Heding & Panning, 1954) P

No data

Elephant Island

600 m

MNHN-IE-2013-

2479

Pentactella leonina (Semper, 1867)

WS 867

Falkland Islands

148-150 m

2016.116-129

Pentactella leonina (Semper, 1867)

WS 84

Falkland Islands

74-75 m

2011.81-90 (23)

Pentactella leonina (Semper, 1867)

WS 85

Falkland Islands

79 m

2011.91-99

Pentactella leonina (Semper, 1867)

D 51

Falkland Islands

115 m

2011.100-101

Pentactella leonina (Semper, 1867)

WS 804

Falkland Islands

143-150 m

2011.102-110

Pentactella leonina (Semper, 1867)

WS 863

Patagonia Shelf

117-121m

2011.111

Pentactella leonina (Semper, 1867)

WS 93

Falkland Islands

130-133 m

2011.112

Pentactella leonina (Semper, 1867)

D 652

Burdwood Bank

169-171m

2011.113

Pentactella leonina (Semper, 1867)

WS 56

Larsen Harbour, South

Georgia

2 m

2016.132-133

Pentactella leonina (Semper, 1867)

D 1909

Falkland Islands

132 m

2016.174

Pentactella leonina (Semper, 1867)

WS 243

Falkland Islands

141-144m

2011.114

Pentactella leonina (Semper, 1867)

WS 81

Falkland Islands

81-82 m

2011.115

Pentactella leonina (Semper, 1867)

WS 80

Falkland Islands

152-156 m

2011.116

Pentactella leonina (Semper, 1867)

WS 865

Patagonia Shelf

126-128 m

2011.117

Pentactella leonina (Semper, 1867)

WS 804

Falkland Islands

143-150 m

2011.118

Pentactella leonina (Semper, 1867)

WS 834

off Patagonia

27-38 m

2011.119

Pentactella leonina (Semper, 1867)

WS 65

Undine Harbour, South

Georgia

1 m (kelp)

2011.120

Pentactella leonina (Semper, 1867)

WS 80

Falkland Islands

152-156 m

2011.121-123

Pentactella leonina (Semper, 1867)

D 52

Falkland Islands

17 m

2011.124

Pentactella leonina (Semper, 1867)

WS 216

Falkland Islands

133-219 m

2011.125

Pentactella leonina (Semper, 1867)

D 51

Falkland Islands

115 m

2011.186-192

Pentactella leonina (Semper, 1867)

WS 804

Falkland Islands

143-150 m

2011.193

Pentactella leonina (Semper, 1867)

WS 56

Larsen Harbour, South

Georgia

2 m

2011.194—195

Pentactella leonina (Semper, 1867)

D 724

Fortesque Bay, Magellan

Strait

0-5 m

2011.184—185

Pentactella leonina (Semper, 1867)

WS 836

Patagonia Shelf

64 m

2011.228

Pentactella leonina (Semper, 1867)

WS 247

Falkland Islands

172 m

2011.229

The Discovery Expedition sea cucumbers (Echinodermata: Holothuroidea)

Taxa (grouped in orders, in alphabetical sequence)

Station

number

Locality

Depth

NHMUK registration

(+ MNHN, ZMUC)

Pentactella leonina (Semper, 1867)

D 53

Port Stanley, Falkland

Islands

0-2 m

2011.230-231

Pentactella leonina (Semper, 1867)

D 55

Port Stanley, Falkland

Islands

10-16 m

2011.232-235

Pentactella leonina (Semper, 1867)

WS 576

Falkland Islands

24-34m

2011.236-237

Pentactella leonina (Semper, 1867)

WS 249

Falkland Islands

166 m

2011.238

Pentactella leonina (Semper, 1867)

WS 848

Patagonia Shelf

115-117 m

2011.239-240

Pentactella leonina (Semper, 1867)

D 53

Port Stanley, Falkland

Islands

0-2 m

2011.241

Pentactella leoninoides (Mortensen, 1925)

D 2215

NZ Antipodes Island

163-210 m

2011.174

Pentactella marionensis (Theel, 1886)

D 1563

Marion Island

101-106m

2011.221-224

Pentactella perrieri (Ekman, 1927)

WS 867

Falkland Islands

148-150 m

2016.99

Pentactella perrieri (Ekman, 1927)

WS 804

Falkland Islands

143-150 m

2016.146

Pentactella perrieri (Ekman, 1927)

WS 840

Falkland Islands

386^163 m

2011.1

Pentactella perrieri (Ekman, 1927)

WS 246

Falkland Islands

208-267 m

2011.2-5

Pentactella perrieri (Ekman, 1927)

D 51

Falkland Islands

115 m

2011.6-9

Pentactella perrieri (Ekman, 1927)

WS 246

Falkland Islands

208-267 m

2011.10-13

Pentactella perrieri (Ekman, 1927)

WS 65

Undine Harbour, South

Georgia

1 m (kelp)

2011.14

Pentactella perrieri (Ekman, 1927)

D 1909

Falkland Islands

132 m

2011.15

Pentactella perrieri (Ekman, 1927)

WS 225

Falkland Islands

161-162m

2011.16

Pentactella perrieri (Ekman, 1927)

D 388

Cape Horn

121 m

2011.17-21

Pentactella perrieri (Ekman, 1927)

WS 243

Falkland Islands

141-144m

2011.22-23

Pentactella perrieri (Ekman, 1927)

WS 228

Falkland Islands

229-236 m

2011.24—26

Pentactella perrieri (Ekman, 1927)

WS 840

Falkland Islands

368^163 m

2011.27

Pentactella perrieri (Ekman, 1927)

WS 228

Falkland Islands

229-236 m

2011.28-30

Pentactella perrieri (Ekman, 1927)

WS 73

Falkland Islands

121-130m

2011.31-33

Pentactella perrieri (Ekman, 1927)

WS 804

Falkland Islands

143-150 m

2011.34-36

Pentactella perrieri (Ekman, 1927)

WS 84

Falkland Islands

74—75 m

2011.37

Pentactella perrieri (Ekman, 1927)

WS 583

Magellan Strait

14-78m

2011.38-40

Pentactella perrieri (Ekman, 1927)

WS 243

Falkland Islands

141-144 m

2011.41-42

Pentactella perrieri (Ekman, 1927)

WS 237

Falkland Islands

150-256 m

2011.43

Pentactella perrieri (Ekman, 1927)

D 1909

Falkland Islands

132 m

2011.44-46

Pentactella perrieri (Ekman, 1927)

WS 88

Falkland Islands

118 m

2011.47-56

Pentactella perrieri (Ekman, 1927)

WS 246

Falkland Islands

208-267 m

2011.57-59

Pentactella perrieri (Ekman, 1927)

WS 825

Falkland Islands

135-144 m

2011.60-69

Pentactella perrieri (Ekman, 1927)

WS 839

Falkland Islands

503-534 m

2011.70

Pentactella perrieri (Ekman, 1927)

WS 804

Falkland Islands

143-150 m

2011.71

Pentactella perrieri (Ekman, 1927)

WS 825

Falkland Islands

135-144 m

2011.72-75

Pentactella perrieri (Ekman, 1927)

WS 93

Falkland Islands

130-133 m

2011.76-77

Pentactella perrieri (Ekman, 1927)

D 141

Cumberland Bay, South

Georgia

17-27 m

2011.78

Pentactella perrieri (Ekman, 1927)

WS 841

Falkland Islands

100-110m

2011.79

Pentactella perrieri (Ekman, 1927)

WS 231

Falkland Islands

159-167 m

2011.80

Pentactella perrieri (Ekman, 1927)

WS 825

Falkland Islands

135-144 m

2011.198-204

Pentamera chiloensis (Ludwig, 1887)

WS 816

Patagonia Shelf

150 m

2011.227

Pentamera chiloensis (Ludwig, 1887)

WS 71

Falkland Islands

80-82 m

2011.225-226

Pseudocnella insolens (Theel, 1886)

MS 82

Saldanha Bay, South Africa

7-14 m

2011.284—293 (10+)

P.M. O’Loughlin, E. Tavancheh & C. Harding

Taxa (grouped in orders, in alphabetical sequence)

Station

number

Locality

Depth

NHMUK registration

(+ MNHN, ZMUC)

Psolicrwc coatsi (Vaney, 1908)

WS 56

Larsen Harbour, South

Georgia

2 m

2011.151-152

Psolicrwc iuvenilesi O’Loughlin & Manjon-Cabeza,

2009b

D 363

South Sandwich Islands

278-329 m

2011.153

Psolicrwc iuvenilesi O’Loughlin & Manjon-Cabeza,

2009b

D 159

South Georgia

160 m

2011.313

Psolidium dorsipes Ludwig, 1887

WS 834

Cape Virgenes, Argentina

27-38 m

2008.3182

Psolidium gaini Vaney, 1914

D 1660

Pennell Bank, Ross Sea

0-351m

2008.3183-3189

Psolidium incubans Ekman, 1925

MS 67

Cumberland Bay, South

Georgia

38 m

2008.3190

Psolidium tenue Mortensen, 1925

D 181

Palmer Archipelago

160-335 m

2008.3191

Psolidium tenue Mortensen, 1925

D 182

Palmer Archipelago

278-500 m

2008.3192

Psolidium tenue Mortensen, 1925

D 187

Palmer Archipelago

0-195 m

2008.3193

Psolidium tenue Mortensen, 1925

D 190

Palmer Archipelago

0-250 m

2008.3194—3196

Psolidium tenue Mortensen, 1925

D 599

Antarctic Peninsula

0-150 m

2008.3197-3198

Psolidium tenue Mortensen, 1925

D 1644

Ross Sea

626 m

2008.3199

Psolidium tenue Mortensen, 1925

D 1649

Ross Sea

695 m

2008.3200-3201

Psolidium tenue Mortensen, 1925

D 1652

Ross Sea

567 m

2008.3202-3205

Psolidium tenue Mortensen, 1925

D 1660

Ross Sea

0-351m

2008.3206-3208

Psolidium tenue Mortensen, 1925

D 2200

Balleny Islands

512-532 m

2008.3209-3218

Psolus antarcticus (Philippi, 1857)

D 55

Port Stanley, Falkland

Islands

10-16 m

2016.69-74

Psolus antarcticus (Philippi, 1857)

D 1909

Burdwood Bank

132 m

2016.75

Psolus antarcticus (Philippi, 1857)

D 1909

Burdwood Bank

132 m

2011.345-354

Psolus antarcticus (Philippi, 1857)

D 1909

Burdwood Bank

132 m

2011.355

Psolus antarcticus (Philippi, 1857)

WS 804

Falkland Islands

143-150 m

2016.76

Psolus antarcticus (Philippi, 1857)

WS 243

Falkland Islands

141-144 m

2016.77

Psolus antarcticus (Philippi, 1857)

WS 825

Falkland Islands

135-144 m

2016.78-80

Psolus antarcticus (Philippi, 1857)

WS 244

Falkland Islands

253-247 m

2016.81

Psolus antarcticus (Philippi, 1857)

D 2200

Balleny Islands

512-532 m

2016.82

Psolus carolineae O’Loughlin & Whitfield, 2010

D 1952

South Shetland Islands

367-383 m

2016.108-110

Psolus dubiosus Ludwig & Heding, 1935

D 1652

Ross Sea

567 m

2016.84—88

Psolus dubiosus Ludwig & Heding, 1935

D 1660

Ross Sea

351 m

2016.89-90

Psolus dubiosus Ludwig & Heding, 1935

D 1652

Ross Sea

567 m

2016.91-92

Psolus dubiosus Ludwig & Heding, 1935

D 474

West of Shag Rocks

199 m

2016.93-98

Psolus figulus Ekman, 1925

MS 67

Cumberland Bay, South

Georgia

38 m

2016.43

Psolus figulus Ekman, 1925

WS 56

Larsen Harbour, South

Georgia

2 m

2016.44-46

Psolus lockhartae O’Loughlin & Whitfield, 2010

WS 840

Falkland Islands

368^63 m

2016.83

Psolusparadubiosus Carriol & Feral, 1985

D 1563

Marion Island

99-113 m

2016.111-112

Psolus patagonicus Ekman, 1925

D 175

South Shetland Islands

200 m

2016.130

Psolus patagonicus Ekman, 1925

D 1957

South Shetland Islands

785-810 m

2016.131

Psolus patagonicus Ekman, 1925

WS 56

East Falklands

10-16 m

2011.376-377

Psolus patagonicus Ekman, 1925

WS 85

Falkland Islands

79 m

2011.356-365

Psolus patagonicus Ekman, 1925

D 1909

Falkland Islands

132 m

2011.369-371

Psolus patagonicus Ekman, 1925

WS 848

Patagonia Shelf

115-117 m

2011.372

Psolus patagonicus Ekman, 1925

D 724

Strait of Magellan

0-5 m

2011.373

The Discovery Expedition sea cucumbers (Echinodermata: Holothuroidea)

Taxa (grouped in orders, in alphabetical sequence)

Station

number

Locality

Depth

NHMUK registration

(+ MNHN, ZMUC)

Psolus patagonicus Ekman, 1925

WS 243

Falkland Islands

141-144 m

2011.374

Psolus patagonicus Ekman, 1925

WS 841

Falkland Islands

100-110m

2011.375

Psolus patagonicus Ekman, 1925

WS 820

Falkland Islands

351-368 m

2011.378

Psolus punctatus Ekman, 1925

D 148

South Georgia

132-148 m

2016.1-24

Psolus punctatus Ekman, 1925

D 474

West of Shag Rocks

199 m

2016.25

Psolus punctatus Ekman, 1925

D 140

South Georgia

122-136 m

2016.26-27

Psolus punctatus Ekman, 1925

D 27

South Georgia

110 m

2016.28

Psolus punctatus Ekman, 1925

D 179

Palmer Archipelago

4-10 m

2016.29-32

Psolus punctatus Ekman, 1925

D 42

Cumberland Bay, South

Georgia

120-204 m

2016.33-39

Psolus punctatus Ekman, 1925

D 190

Palmer Archipelago

130 m

2016.40^-2

Staurocucumis liouvillei (Vaney, 1914)

D 456

Ross Sea

40-45 m

2016.137-139

Staurocucumis liouvillei (Vaney, 1914)

WS 225

Falkland Islands

161-162m

2016.155

Staurocucumis liouvillei (Vaney, 1914)

D 140

South Georgia

122-136 m

2011.159-161

Staurocucumis liouvillei (Vaney, 1914)

D 366

South Sandwich Islands

77-152 m

2011.154-158

Staurocucumis liouvillei (Vaney, 1914)

D 123

South Georgia

230-250 m

2011.340

Trachythyone bouvetensis (Ludwig & Heding, 1935)

D 27

Cumberland Bay, South

Georgia

110 m

2016.142

Trachythyone bouvetensis (Ludwig & Heding, 1935)

D 144

South Georgia

155-178 m

2011.175

Trachythyone bouvetensis (Ludwig & Heding, 1935)

D 160

Shag Rocks

0-180 m

2011.176-177

Trachythyone bouvetensis (Ludwig & Heding, 1935)

D 1652

Ross Sea

567 m

2011.178

Trachythyone bouvetensis (Ludwig & Heding, 1935)

D 170

South Shetland Islands

342 m

2011.179-180

Trachythyone bouvetensis (Ludwig & Heding, 1935)

D 1660

Ross Sea

351 m

2011.181-182

Trachythyone bouvetensis (Ludwig & Heding, 1935)

D 1658

Ross Sea

520 m

2011.183

Trachythyone bouvetensis (Ludwig & Heding, 1935)

WS 71

Falkland Islands

80-82 m

2011.337-338

Trachythyone bouvetensis (Ludwig & Heding, 1935)

D 42

Cumberland Bay, South

Georgia

120-204 m

2011.339

Trachythyone parva (Ludwig, 1875)

WS 795

Falkland Islands

157-161m

2016.103

Thy one aurea (Quoy & Gaimard, 1834)

MS 82

Saldanha Bay, South Africa

7-14 m

2011.294—296

Elasipodida

Peniagone vignoni Herouard, 1901

D 181

Palmer Archipelago

160-335 m

2011.311-312

Rhipidothuria racovitzae Herouard, 1901

D 181

Palmer Archipelago

160-335 m

2011.298-307(20)

Rhipidothuria racovitzae Herouard, 1901

D 1952

South Shetland Islands

367-383 m

2011.308-310

Synaptida (sensu Smirnov 2012)

Chiridota pisanii Ludwig, 1887

WS 750

off Patagonia

95 m

2010.110

Chiridota pisanii Ludwig, 1887

WS 388

Cape Horn

121 m

2010.111

Paradota weddellensis Gutt, 1990

WS 217

Falkland Islands

146 m

2011.242

Paradota weddellensis Gutt, 1990

D 600

South Shetland Islands

501-527 m

2011.243

Scoliorhapis massini O’Loughlin & VandenSpiegel,

2010

WS 756

Falkland Islands

119 m

2010.105-109

Scoliorhapis massini O’Loughlin & VandenSpiegel,

2010 (“probably”)

MS 67

Cumberland Bay, South

Georgia

38 m

2010.112-113

Sigmodota contorta (Ludwig, 1875)

WS 816

Patagonia Shelf

150 m

2010.55-62

Sigmodota contorta (Ludwig, 1875)

WS 88

Falkland Islands

118 m

2010.63-68

Sigmodota contorta (Ludwig, 1875)

WS 84

Falkland Islands

75 m

2010.69-70

Sigmodota contorta (Ludwig, 1875)

WS 773

Falkland Islands

296 m

2010.71-74

Sigmodota contorta (Ludwig, 1875)

WS 82

Falkland Islands

140-144 m

2010.75-84

P.M. O’Loughlin, E. Tavancheh & C. Harding

Taxa (grouped in orders, in alphabetical sequence)

Station

number

Locality

Depth

NHMUK registration

(+ MNHN, ZMUC)

Sigmodota contorta (Ludwig, 1875)

WS 25

Undine Harbour, South

Georgia

18-27 m

2010.85-94

Sigmodota contorta (Ludwig, 1875)

WS 56

Larsen Harbour, South

Georgia

2010.95-96

Sigmodota contorta (Ludwig, 1875)

D 39

Cumberland Bay, South

Georgia

179-235 m

2010.97-98

Sigmodota contorta (Ludwig, 1875)

D 126

South Georgia

0-100 m

2010.99

Sigmodota contorta (Ludwig, 1875)

WS 62

Wilson Harbour, South

Georgia

15^5 m

2010.100-103

Sigmodota contorta (Ludwig, 1875)

D 1941

South Georgia

22-25 m

2010.104

Sigmodota contorta (Ludwig, 1875)

D 1562

Marion Island

88-104m

2016.47-68

Sigmodota contorta (Ludwig, 1875)

WS 83

Falkland Islands

129-137 m

2011.138

Sigmodota contorta (Ludwig, 1875)

WS 804

Falkland Islands

143-150 m

2011.205-211

Sigmodota contorta (Ludwig, 1875)

WS 228

Falkland Islands

229-236 m

2011.212-220

Sigmodota contorta (Ludwig, 1875)

WS 869

Patagonia Shelf

187 m

2011.244—253 (50+)

Sigmodota contorta (Ludwig, 1875)

D 45

South Georgia

238-270 m

2011.254-263 (20+)

Sigmodota contorta (Ludwig, 1875)

D 145

South Georgia

26-35 m

2011.264—273 (10+)

Sigmodota contorta (Ludwig, 1875)

WS 838

Patagonia Shelf

149-159 m

2011.274-283(10+)

4 Sigmodota magnibacula (Massin & Heterier, 2004)

2016.144

taeniogyrinid species (very poor condition)

WS 182

Palmer Archipelago

278-500 m

2010.114

^ee note 1 with Table 2 below.

2 See note 2 with Table 2 below.

3 Holotype specimen is thought to be lost. See note 3 with Table 2 below.

4 Specimen and locality source not recognized. May not be Discovery Expedition. Handwritten label with: St. 105,13/2/’31, 163 m.

Etymology. Named wesafrica for the geographical occurrence

of the species on the West Africa coast.

Remarks. All the type specimens have dried during

preservation, and the calcareous ring and ossicles are partly at

least eroded. The ring does not have a recognizable outline in

any specimen that was dissected. The form and number of the

tentacles are difficult to observe. The sub-pentagonal form,

complete body cover of imbricating scales with free ends

pointing posteriorly, 10 short lobed tentacles, radial series of

tube feet, and absence of cups or tables distinguish the new

genus and species.

Previous publications with Discovery Expedition

Holothuroidea (included in Table 1 and listed in Tables 2, 3,

4).

1. Albert Panning (in Heding & Panning 1954) recorded that

Svend Heding died (in 1949) before their manuscript was

completed, and before Heding was able to complete any

Discovery Expedition Report. Panning acknowledged that

descriptions of the new genera and species Clarkiella Heding,

1954, Clarkiella discoveryi Heding, 1954, Parathyonidium

Heding, 1954 and Parathyonidium incertum Heding, 1954

were from the notes of Heding. In collaboration with Elizabeth

Deichmann (at the MCZ at Harvard University at the time)

Panning included these new taxa in the comprehensive paper

Heding & Panning 1954. Panning (in Heding & Panning 1954)

recorded that the work on the Discovery material was taken

over by Dr. Deichmann. This was not completed, and no

Discovery Report has been published.

O’Loughlin (2009), in a paper on the BANZARE

holothuroids, discussed Clarkiella Heding, and referred a new

holothuroid species from the Kerguelen Islands and Tasmania

to Heding’s genus: Clarkiella deichmannae O’Loughlin,

2009.

O’Loughlin et al. (2009a), in reporting some observations

of reproductive strategies of dendrochirotid species, referred

to an as then undescribed species of Parathyonidium Heding

from Eastern Antarctica that exhibited brood-protection in the

coelom. Subsequently O’Loughlin et al. (2014) included an

illustrated revision of genus Parathyonidium Heding and

species Parathyonidium incertum Heding. Their revision was

based on the original description and on three paratypes from

the South Shetland Islands. Additional specimens were

recognized from South Georgia, from the Antarctic Peninsula,

and off Enderby Land in Eastern Antarctica. The specimen

referred to Parathyonidium by O’Loughlin et al. (2009a) was

identified as P. incertum. ZMUC records (see note 3 under

Table 2) indicate that the holotype for Parathyonidium

incertum “must probably be considered lost”. O’Loughlin et

al. (2014) listed three lots of paratypes that are held respectively

in Copenhagen (ZMUC), Paris (MNHN) and London

The Discovery Expedition sea cucumbers (Echinodermata: Holothuroidea)

Figure 5. a & b, photos of preserved holotype of Cucusquama wesafrica O’Loughlin gen. & sp. nov. from Angola (oral end left; axial length 13

mm; NHMUK 2016.148); c, photo into mouth cavity of a preserved paratype from Gambon (French Congo) (arrow pointing to one black tentacle;

NHMUK 2016.149).

(NHMUK). Details are provided in Tables 1 & 2. O’Loughlin

et al. (2014) reported the holotype locality for Parathyonidium

incertum as Shag Rocks near South Georgia, based on Heding

& Panning (1954). We judge here that this was a mistake (see

note 2 with Table 2 below).

2. O’Loughlin & Ahearn (2008) reported 13 lots of Discovery

Expedition holothuroids that represent four species of Psolidium

Ludwig, 1875. Details are provided in Tables 1 and 3.

3. O’Loughlin & VandenSpiegel (2010) reported 16 lots of

Discovery Expedition Synaptida (as Apodida) holothuroids

that represented 3 species. Details are provided in Tables 1

and 4.

Systematic notes on Discovery holothuroid taxa.

1. Echinopsolus Gutt, 1990.

Bohn & Hess (2014) reassigned a group of Antarctic

cucumariid species to genus Echinopsolus Gutt, 1990, based

on their shared and unique set of morphological characters

related to their reproductive mode. The group comprised:

Echinopsolus acanthocola Gutt, 1990; E. acutus (Massin,

1992); E. charcoti (Vaney, 1906); E. koehleri (Vaney, 1914); E.

mollis (Ludwig & Heding, 1935); E. parvipes (Massin, 1992);

E. splendidus (Gutt, 1990). In the same paper Bohn & Hess

(2014) reassigned genus Echinopsolus to family Cucumariidae.

P.M. O’Loughlin, E. Tavancheh & C. Harding

Table 2. Type specimens of Discovery Expedition Holothuroidea published in Heding & Panning 1954.

Taxon

Type status

Station

collected

Locality

collected

Depth

Date

collected

Institution

lodged

Registration

Clarkiella discoveryi Heding

(in Heding & Panning, 1954)

'Holotype

D 474

W of Shag

Rocks

South Georgia

199 m

19 Nov

1930

ZMUC

HOL-000064

Clarkiella discoveryi Heding,

1954

'Paratype

D 474

W of Shag

Rocks

South Georgia

199 m

19 Nov

1930

ZMUC

HOL-000247

Parathyonidium incertum

Heding

(in Heding & Panning, 1954)

2 'Holotype

3 Lost

specimen

3 HOL-000093

Parathyonidium incertum

Heding, 1954

Paratypes (3)

D 170

Clarence Island

S Shetland

Islands

342 m

23 Feb 1927

ZMUC

HOL-000300

Parathyonidium incertum

Heding, 1954

Paratypes (3)

D 170

Clarence Island

342 m

23 Feb 1927

NHMUK

2011.171-173

Parathyonidium incertum

Heding, 1954

Partypes (2)

No record

Elephant Island

600 m

No record

MNHN

MNHN-

IE-2013-2479

'No Discovery station data were reported for Clarkiella discoveryi with the description of the new taxa in Heding & Panning

1954, but registered and labelled holotype and paratype (one) specimens are in the ZMUC with type status, station number and

collection data (see Table 2 above with station data from the labels with the types in the ZMUC). Both type specimens were

collected from the same type locality, station D474.

2 The station data reported for the type for Parathyonidium incertum in Heding & Panning (1954) is station D474. We judge that

this may be a mistake since it is the type locality on the labels for Clarkiella discoveryi. The holotype specimen is assumed to

be lost as no “holotype” has been found. But there are paratypes from station D170, and a note on the label with them translated

by Tom Schioette in 2013 reads: “Does the identification with them include also the large specimens? Heding’s serial number

234-236 st. 170”. With some reservation we judge that the holotype was most probably also from the paratype station D170, and

not station D474 as published in Heding & Panning (1954).

3 Note by Tom Schioette in 2013: “The holotype of Parathyonidium incertum, which should probably have been (or perhaps was)

returned with the “Discovery” material after Heding’s death, was later given the ZMUC number HOL-93 in absentia. It must

probably be considered lost, since later workers on the material have not succeeded in finding it”.

. d mm o

Figure 6. Photos of eroding ossicles from the mid-body wall of Cucusquama wesafrica O’Loughlin gen. & sp. nov. a, large single-layered

perforated plates (scales) from the holotype (up to 600 fim long; NHMUK 2016.148); b, body wall and tube foot small perforated plates from a

paratype (up to 168 fim long; NHMUK 2016.150).

The Discovery Expedition sea cucumbers (Echinodermata: Holothuroidea)

Table 3. Discovery Expedition Holothuroidea published in O’Loughlin & Ahearn 2008.

Taxon

Station

collected

Locality collected

Depth

Institution

lodged

Registration

Psolidium dorsipes Ludwig, 1887

WS 834

Cape Virgenes

Southern Argentina

27-38 m

NHMUK

2008.3182

Psolidium gaini Vaney, 1914

D 1660

Pennell Bank

Ross Sea

0-351m

NHMUK

2008.3183-3189

Psolidium incubans Ekman, 1925

MS 67

South Georgia

38 m

NHMUK

2008.3190

Psolidium tenue Mortensen, 1925

D 181

Palmer Archipelago

Antarctica

160-335 m

NHMUK

2008.3191

Psolidium tenue Mortensen, 1925

D 182

Palmer Archipelago

Antarctica

278-500 m

NHMUK

2008.3192

Psolidium tenue Mortensen, 1925

D 187

Palmer Archipelago

Antarctica

0-195 m

NHMUK

2008.3193

Psolidium tenue Mortensen, 1925

D 190

Palmer Archipelago Antarctica

0-250 m

NHMUK

2008.3194—3196

Psolidium tenue Mortensen, 1925

D 599

W of Adelaide Island

Antarctic Peninsula

0-150 m

NHMUK

2008.3197-3198

Psolidium tenue Mortensen, 1925

D 1644

Ross Sea

626 m

NHMUK

2008.3199

Psolidium tenue Mortensen, 1925

D 1649

Ross Sea

695 m

NHMUK

2008.3200-3201

Psolidium tenue Mortensen, 1925

D 1652

Ross Sea

0-500 m

NHMUK

2008.3202-3205

Psolidium tenue Mortensen, 1925

D 1660

Ross Sea

0-351m

NHMUK

2008.3206-3208

Psolidium tenue Mortensen, 1925

D 2200

Balleny Island

Antarctica

512-532 m

NHMUK

2008.3209-3218

Table 4. Discovery Expedition Holothuroidea published in O’Loughlin & VandenSpiegel 2010.

Taxon

Station

collected

Locality collected

Depth

Institution

lodged

Registration

number

Sigmodota contorta (Ludwig, 1875)

WS 816

Falkland Islands

150 m

NHMUK

2010.55-62

Sigmodota contorta (Ludwig, 1875)

WS 88

Falkland Islands

118 m

NHMUK

2010.63-68

Sigmodota contorta (Ludwig, 1875)

WS 84

Falkland Islands

75 m

NHMUK

2010.69-70

Sigmodota contorta (Ludwig, 1875)

WS 773

Falkland Islands

296 m

NHMUK

2010.71-74

1 Sigmodota contorta (Ludwig, 1875)

WS 82

Falkland Islands

140-144 m

NHMUK

2010.75-84

Sigmodota contorta (Ludwig, 1875)

WS 25

South Georgia

18-27 m

NHMUK

2010.85-94

Sigmodota contorta (Ludwig, 1875)

WS 56

South Georgia

2 m

NHMUK

2010.95-96

Sigmodota contorta (Ludwig, 1875)

D 39

South Georgia

179-235 m

NHMUK

2010.97-98

Sigmodota contorta (Ludwig, 1875)

D126

South Georgia

0-100 m

NHMUK

2010.99

Sigmodota contorta (Ludwig, 1875)

WS 62

South Georgia

15-45 m

NHMUK

2010.100-103

Sigmodota contorta (Ludwig, 1875)

D 1941

South Georgia

22-55 m

NHMUK

2010.104

Scoliorhapis massini

O’Loughlin & VandenSpiegel, 2010

WS 756

Falkland Islands

119 m

NHMUK

2010.105-109

Chiridota pisanii Ludwig, 1887

WS 750

Patagonia

95 m

NHMUK

2010.110

Chiridota pisanii Ludwig, 1887

WS 388

Cape Horn

South America

121 m

NHMUK

2010.111

^ee Systematic note 6 below. This entry has been changed from the O’Loughlin & VandenSpiegel 2010 paper.

P.M. O’Loughlin, E. Tavancheh & C. Harding

O’Loughlin et al. (2009a) discussed the “ Cucumaria

georgiana (Lampert, 1886) group” of Antarctic species that

was created by Gutt (1988), and followed by Massin (1992).

O’Loughlin et al. (2009a) listed 11 species in this “group”:

Cucumaria acuta Massin, 1992; Cucumaria analis Vaney,

1908; Cucumaria aspera Vaney, 1908; Cucumaria attenuata

Vaney, 1906; Cucumaria georgiana (Lampert, 1886);

Cucumaria joubini Vaney, 1914; Cucumaria lateralis Vaney,

1906; Cucumaria perfida Vaney, 1908; Cucumaria periprocta

Vaney, 1908; Cucumaria secunda Vaney, 1908; Cucumaria

vaneyi Cherbonnier, 1949. Bohn & Hess (2014) also discussed

this “group”, and we agree that the systematic status of the

species in this group requires resolution. Foundational to this

systematic resolution must be an establishment of the

systematic status of Cucumaria georgiana (Lampert, 1886).

Bohn & Hess (2014) did not assign the “group” to Echinopsolus.

We have assigned some Discovery Expedition lots to this

“group”. Based on the general similarity of their reproductive

morphological features with those of the Echinopsolus species

we have also assigned this “group” to Echinopsolus.

Bohn & Hess (2014) were not able to confirm the

systematic status of Echinopsolus excretiospinosus Massin,

2010, but noted that no brood pouches were reported and the

ventral tentacle pair were apparently not smaller than the

other tentacles.

COl genetic data (Gustav Paulay pers. comm.-, see

phylogenetic tree in O’Loughlin et al. 2011) support a generic

clade that includes Echinopsolus acanthocola (with apparently

two or three cryptic species with geographic congruence), the

“georgiana group” (with apparently two or three cryptic

species with geographic congruence), and the reassigned

Echinopsolus mollis (apparently two or three cryptic species

with geographic congruence). Generic data thus support in

part the work of Bohn & Hess (2014). We note that these

species also have mid-body dorsal papillae or tube feet, and

lack cup (bowl) ossicles in the body wall.

But COl genetic data (Gustav Paulay pers. comm.-, see

phylogenetic tree in O’Loughlin et al. 2011) support a generic

clade for Psolus koehleri and Psolus charcoti that is separate

from the Echinopsolus clade and do not support the

reassignment of these two species to Echinopsolus. We note

that these two species lack mid-body dorsal tube feet or

papillae, and do have cup (bowl) ossicles in the body wall.

Genetic data to date do not support their assignment to a

Psolus Oken, 1815 clade. We leave these two species in their

current reassignment to Echinopsolus until a necessary

reassessment of dendrochirotid generic assignments is

supported by additional genetic data.

We do not have a COl sequence for the recently

reassigned Echinopsolus splendidus. This species lacks dorsal

and lateral tube feet / papillae, but also lacks cups / bowls in

the body wall. It falls morphologically into neither

Echinopsolus genetic /generic clade. We judge that it will

probably fall into another generic clade but in the absence of

supportive genetic data we do not change the current

reassignment to Echinopsolus.

2. Ocnus capensis (Theel, 1886).

We have identified a single Discovery Expedition specimen

from the sub-littoral of Saldanha Bay in south-west South

Africa as Ocnus capensis (Theel, 1886) (MS 82, off

Salamander Point, 7-14 m, 6 Sept 1926, NHMUK 2016.143).

We based our determination on the description and illustration

by Theel (1886) of the three type specimens collected from

179-274 meters off Cape Town in South Africa. Saldanha

Bay is close to the type locality for this species. Based on our

laboratory notes and sketches, Frank Rowe {pers. comm)

judged that the species is Ocnus capensis, but thought that the

species would be better assigned to Pseudocnus Panning,

1949. Rowe judged that genus Ocnus Forbes & Goodsir, 1839

(in Forbes, 1841) is restricted to the Mediterranean and north

European shore, and that genus Pentacta Goldfuss, 1820 is a

monotypic endemic South Africa genus. Thandar (1991)

described and illustrated and discussed Ocnus capensis, and

Ahmed Thandar (pers. comm.) expressed some doubt about

our identification. He considered the species to be a deep

water one. We acknowledge that there is thus some doubt

about our identification.

3. Pentactella Verrill, 1876.

Many Discovery Expedition lots have been identified as

species of Pentactella Verrill, 1876. Based on morphological

characters and distribution consideration, and with the support

of some genetic data, O’Loughlin et al. (2014) reassigned

numbers of species of Pseudocnus Panning, 1949 to a new

genus Laevocnus O’Loughlin (in O’Loughlin et al. 2014).

Immediately after publication the authors recognized that the

type species for the new genus Laevocnus was the type species

for the monotypic Pentactella Verrill, 1876. Laevocnus is an

objective junior synonym of Pentactella. A detailed systematic

history of genus Pentactella, and the assigned species, is

provided by O’Loughlin et al. (2015).

4. Psolus dubiosus Ludwig & Heding, 1935

COl phylogenetic data (Gustav Paulay pers. comm.) strongly

support a synonymy for Psolus arnaudi Cherbonnier, 1974

and Psolus cherbonnieri Carriol & Feral, 1985 with Psolus

dubiosus Ludwig & Heding, 1935. For Discovery Expedition

specimens we have not attempted to distinguish the former

from Psolus dubiosus.

5. Psolus lockhartae O’Loughlin & Whitfield, 2010.

We have identified a single specimen from deep water off the

Falkland Islands as Psolus lockhartae O’Loughlin &

Whitfield, 2010 (WS 840, S of Falkland Islands, 368-463 m,

6 Feb 1932, NHMUK 2016.83). The distribution of Psolus

lockhartae was given by O’Loughlin & Whitfield (2010) as

Birdwood Bank, South Georgia, South Shetlands and South

Orkneys (211-2897 m). The 12 mm long specimen is smaller

than the types (up to 20 mm long). The ossicle complement is

the same, and the form of the ossicles is similar but the

ossicles in the types are larger. We thus have some reservation

over our determination.

The Discovery Expedition sea cucumbers (Echinodermata: Holothuroidea)

6. Sigmodota contorta (Ludwig, 1875).

O’Loughlin & VandenSpiegel (2010) published the

determinations of numbers of Discovery Expedition synaptid

(as apodid) holothuroids (see Table 4 above). They reported 10

specimens of Sigmodota contorta (Ludwig, 1875) (NHMUK

2010.75-84) from Marine Station 82 (Saldanha Bay). The

location of Sladanha Bay in South Africa was not noticed, and

the locality was mistakenly given as the Falkland Islands.

There have been no other reports of Sigmodota contorta from

South Africa, and this report for Saldanha Bay is now judged

to be a mistake. There is also an RRS William Scoresby station

82 and this is now judged to be the source of the specimens.

This station WS 82 was off the Falkland Islands at 140-144 m.

Acknowledgments

We acknowledge with appreciation those who have studied this

historic collection of sea cucumbers: the late Svend Heding,

Albert Panning, Elizabeth Deichmann and Cynthia Ahearn;

and recent assistants Melanie Mackenzie and Emily Whitfield.

We are grateful for the variety of assistance generously given

by: the late Cynthia Ahearn (USNM, forwarding of the

collection to Museum Victoria); Ben Boonen (confrere of

O’Loughlin, photoshop and format of the figures); Andrew

Cabrinovic (NHMUK, facilitation of registrations of

specimens); David Pawson (Emeritus Senior Scientist, National

Museum of Natural History, Smithsonian Institution, provision

of photos and information on the history of the collection);

Frank Rowe (Research Associate of the Australian Museum,

dialogue on systematic issues); Tom Schioette (ZMUC,

provision of a photo and dialogue on types in the ZMUC);

Ahmed Thandar (University of KwaZulu-Natal, dialogue on

systematic issues). We are grateful to Tom Schioette (ZMUC

pers. comm) for the restoration of an honourable reputation to

Svend Heding who was maligned within our field and

elsewhere. He wrote of Heding: “During the war he was used a

lot by the Germans as an example of scientific cooperation of

foreign researchers with German ones, and that led to some

isolation for him in Denmark, although he was far from being

a sympathizer with German politics (on the contrary he was

among those who helped the Danish Jews to escape to Sweden).

People who knew him said he was a kind and helpful man.” We

are most appreciative of the critical and helpful reviews of our

manuscript generously provided by Melanie Mackenzie

(NMV) and David Pawson (USNM).

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Memoirs of Museum Victoria 75:71-82 (2016) Published 2016

1447-2554 (On-line)

http://museumvictoria.com.au/about/books-and-journals/journals/memoirs-of-museum-victoria/

Suction feeding preceded filtering in baleen whale evolution

Felix G. Marx 1 " 3 *, David P. Hocking 1,2 , Travis Park 1,2 , Tim Ziegler 2 , Alistair R. Evans 1,2 and

Erich M.G. Fitzgerald 1,2,4,5

1 School of Biological Sciences, Monash University, 18 Innovation Walk, Clayton, Victoria 3800, Australia.

2 Geosciences, Museum Victoria, Melbourne, Australia.

3 Directorate of Earth and History of Life, Royal Belgian Institute of Natural Sciences, Brussels, Belgium.

4 National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.

5 Department of Life Sciences, Natural History Museum, London, UK.

* To whom correspondence should be addressed. E-mail: felix.marx@monash.edu

Abstract Marx F.G., Hocking D.R, Park T., Ziegler T., Evans A.R. and Fitzgerald, E.M.G. 2016. Suction feeding preceded filtering

in baleen whale evolution. Memoirs of Museum Victoria 75: 71-82.

The origin of baleen, the key adaptation of modern whales (Mysticeti), marks a profound yet poorly understood

transition in vertebrate evolution, triggering the rise of the largest animals on Earth. Baleen is thought to have appeared in

archaic tooth-bearing mysticetes during a transitional phase that combined raptorial feeding with incipient bulk filtering.

Here we show that tooth wear in a new Late Oligocene mysticete belonging to the putatively transitional family Aetiocetidae

is inconsistent with the presence of baleen, and instead indicative of suction feeding. Our findings suggest that baleen arose

much closer to the origin of toothless mysticete whales than previously thought. In addition, they suggest an entirely new

evolutionary scenario in which the transition from raptorial to baleen-assisted filter feeding was mediated by suction,

thereby avoiding the problem of functional interference between teeth and the baleen rack.

Keywords Mysticeti; baleen whale; filter feeding; suction feeding; tooth wear; Aetiocetidae

Introduction

Baleen whales (Mysticeti) are the largest animals on Earth

and owe their success to baleen, a unique feeding structure

allowing them to filter vast quantities of small prey from

seawater (Pivorunas, 1979; Werth, 2000b). The baleen

apparatus of extant mysticetes consists of a series of keratinous

plates suspended from the upper jaw, traditionally thought to

be derived from the horny palatal ridges of extant artiodactyls

(Werth, 2000b). More recent anatomical work, however, has

shown that the basal tissue giving rise to baleen is innervated

by the superior alveolar nerves, and is thus more likely

homologous with the gingiva (Sawamura, 2008).

Baleen rarely fossilises (Esperante et al., 2008; Gioncada

et al., 2016), but is thought to have originated early in mysticete

evolution, during a transitional phase combining tooth-based

raptorial feeding and baleen-assisted filtering (Demere and

Berta, 2008; Demere et al., 2008). This transition is seemingly

exemplified by the Aetiocetidae - a mostly Oligocene (34-23

Ma) family of archaic mysticetes which retained functional

teeth alongside features commonly associated with filter

feeding (Demere et al., 2008). The underlying drivers,

mechanics and accuracy of this scenario, however, remain

contentious (Fitzgerald, 2010; Marx et al., 2015). Here we

show that a new Late Oligocene aetiocetid fossil from

Washington, USA, has a highly distinctive tooth wear pattern

that is inconsistent with the presence of baleen, suggesting that

this key mysticete adaptation emerged later and much closer to

the origin of modern whales than previously thought. Our new

fossil displays functional adaptations for suction feeding

rather than filtering, casting doubt on the accepted

ecomorphological context of chaeomysticete evolution. Based

on this new information, we re-examine previous arguments

in favour of a direct transition from raptorial to filter feeding,

and propose an alternative model of baleen evolution more

consistent with available evidence both from extant taxa and

the fossil record.

Material and Methods

Except for the right p3, the teeth were found encased in soft

sediment and washed out using water, with no mechanical or

chemical preparation. The remainder of the specimen was

prepared mechanically and using 10% acetic acid. All parts of

the specimen in figs 1-3 were coated with ammonium chloride

prior to photography. Where appropriate, photographs of the

specimen were taken at varying foci and digitally stacked in

Photoshop CS6. To visualise the gross wear features further,

we scanned the two best-preserved teeth via micro-computed

tomography using a Zeiss Xradia 520 Versa (Oberkochen,

F.G. Marx, DP. Hocking, T. Park, T. Ziegler, A.R. Evans & E.M.G. Fitzgerald

Germany) at the Monash University X-ray Microscopy Facility

for Imaging Geomaterials (XMFIG). The specimens were

scanned with a source voltage of 140 kV and current of 70 pA,

an exposure time of 2 seconds per image and a pixel size of

12.7 pm. 3D surface models were generated in Avizo v9.0.0

(Visualization Science Group) and are available as

supplementary 3D figures (figs S1-S2).

Institutional abbreviations

AMP, Ashoro Museum of Paleontology, Ashoro, Hokkaido,

Japan; LACM, Natural History Museum of Los Angeles

County, Los Angeles, USA; NMV, Museum Victoria,

Melbourne, Australia; UCMP, University of California

Museum of Paleontology, Berkeley, USA; USNM, National

Museum of Natural History, Smithsonian Institution,

Washington, DC, USA; UWBM, Burke Museum of Natural

History and Culture, University of Washington, Seattle, USA.

Results

Description of NMV P252567

The new fossil specimen (NMV P252567) comes from the

upper part of the Pysht Formation (Clallam County,

Washington State, USA; Late Oligocene) (Prothero et al.,

2001), and comprises much of the cranium, both mandibles,

and postcranial elements. Overall, the morphology of the

skull is intermediate between that of Aetiocetus and Fucaia

(Barnes et al., 1995). A detailed systematic analysis of NMV

P252567 is currently under preparation, but it is confidently

assigned to Aetiocetidae based on the presence of (i) an

enlarged lacrimal incising into the lateral border of the

ascending process of the maxilla (Demere and Berta, 2008;

Marx et al., 2015); (ii) a laterally expanded premaxilla

overhanging the adjacent portion of the maxilla (Barnes et al.,

1995; Fitzgerald, 2010; Geisler and Sanders, 2003; Marx,

2011); (iii) a proportionally large, anterolaterally directed

orbit (also present in Mammalodontidae) (Marx, 2011); (iv) a

(presumably ligamentous) mandibular symphysis with

attendant symphyseal groove (also present in chaeomysticetes)

(Fitzgerald, 2012); (v) gracile cheek teeth with fused roots

(Demere and Berta, 2008; Marx et al., 2015); and (vi)

lingually ornamented tooth crowns (Demere and Berta, 2008;

Marx et al., 2015) (figs 1-3).

Based on the right mandible, NMV P252567 has 11 lower

teeth, similar to archaeocetes, mammalodontids and Fucaia

goedertorum (Barnes et al., 1995; Fitzgerald, 2010; Uhen, 2004).

There are at least nine preserved teeth, including: a left (II or 12)

and a right upper incisor (12 or 13); the left upper canine or PI;

the in situ roots of right p3; and five double-rooted postcanines,

including at least one upper and one lower (figs 1-3, S1-S2). The

left incisor has a broken apex, but otherwise displays intact labial

and lingual enamel surfaces with no obvious abrasive wear (fig.

3A). The right incisor is abraded along two thirds of its lingual

surface, but intact labially (fig. 3B).

All of the remaining teeth are heavily abraded with the

consequent loss of all lingual enamel, except for a thin band

along the base of the crown that was presumably located below

the gum line (fig. 2). Between this basal band and the apex, the

lingual surface of each crown is deeply excavated and polished.

Where preserved, the centre of the polished surface bears

several deep, horizontal striations, the edges of which are

themselves polished and rounded (fig. 2). Anteriorly and

posteriorly, the abraded surface wraps around the crown,

resulting in an hourglass-shaped labial wear pattern. The

extent of labial wear varies, but in at least one tooth all of the

enamel has been removed except for a centrally located,

vertical strip (fig. 2B). The most heavily worn teeth, which are

likely also the most posterior, have lost most of their crowns

and are reduced to a basal band of enamel and a lingually

excavated, low remnant of dentine (figs 2C, 3C).

Comparisons with Aetiocetidae and other marine mammals

The pattern and intensity of tooth wear in NMV P252567 is

unique among Aetiocetidae. Besides the present material,

tooth wear has been described in some detail for three

aetiocetids, namely, Aetiocetus cotylalveus, A. weltoni and

Fucaia buelli (Demere and Berta, 2008; Emlong, 1966; Marx

et al., 2015). In addition, teeth are preserved but have not been

properly figured in A. polydentatus and Morawanocetus

yabukii (Barnes et al., 1995). The enamel covering the crowns

in all of these species lacks the heavy abrasion characteristic

of NMV P252567. Several of the premolars and molars in the

holotypes of A. cotylalveus (USNM 25210) and A. weltoni

(UCMP 122900) instead show attritional wear, which has

removed much or all of the accessory denticles (Demere and

Berta, 2008; Emlong, 1966). In addition, relatively minor

apical abrasion is evident along at least the anterior portion of

the tooth row in A. weltoni, and on both the premolars and

molars of A. cotylalveus.

Tooth wear in A. polydentatus has not been described in

detail, but (presumably attritional) wear facets seemingly

occur at least on the posteriormost mandibular teeth (Demere

and Berta, 2008). Both attrition and apical abrasion also occur

in Fucaia buelli, with extensive attritional wear facets

occurring on the lingual surfaces of the upper premolars and

molars of the type specimen (UWBM 84024; Marx et al.,

2015). Too little is known about Morawanocetus to be sure

about wear patterns in this species. Nevertheless, based on

photographs, at least one of the posterior molars preserved

with the holotype (AMP 01) displays strong apical and,

possibly, attritional wear.

In general, the dental wear of NMV P252567 most closely

resembles that of the bizarre-looking archaic mysticete

Mammalodon colliveri and the extant walrus, Odobenus

rosmarus, both of which show lingual abrasion and

(microscopic) striations, and are considered to be benthic

suction feeders (Fitzgerald, 2010; Gordon, 1984). Unlike

NMV P252567, however, M. colliveri has small, peg-like

incisors displaying heavy abrasion, and its dentition is

generally even more heavily worn (Fitzgerald, 2010). Other

marine mammals known to show heavy dental wear include

the killer whale Orcinus orca, the beluga Delphinapterus

leucas, the porpoises Phocoena phocoena and Semirostrum

ceruttii, and the archaic beaked whale Ninoziphiusplatyrostris.

However, in orcas such wear generally consists of severe

apical abrasion, possibly as a result of preying on sharks (Ford

Suction feeding preceded filtering in baleen whale evolution

lacrimal

incising

into maxilla

premaxilla

overhanging

maxilla

A^la

large orbit

embrasure

pits

Figure 1. Diagnostic characteristics identifying NMV P252567 as an aetiocetid. A, explanatory line drawing of the skull; B, photograph of the

skull (left) and mandible (right), both in dorsal view.

et al., 2011), while in belugas direct tooth-on-tooth occlusion

results in a predominance of attrition (Fitzgerald, 2010;

Struthers, 1895). By contrast, heavy wear in phocoenids and

N. platyrostris may primarily reflect benthic foraging and the

frequent ingestion of abrasive sediment (Lambert et al., 2013;

Racicot et al., 2014), although more recent studies suggest that

stem ziphiids may have foraged on epipelagic prey (Lambert

et al., 2015).

Discussion and Conclusions

Feeding strategy ofNMVP252567

The distinctive wear pattern of NMV P252567 provides

insights into its likely feeding method. In particular, the

pronounced lingual excavation of the crowns and attendant

striations suggest that the insides of the teeth were subject to

strong abrasive forces, such as repeated anteroposterior

F.G. Marx, DP. Hocking, T. Park, T. Ziegler, A.R. Evans & E.M.G. Fitzgerald

worn

surface

worn

surface

horizontal

striations

unworn

enamel

horizontal

striations

worn

surface

unworn

enamel

carina

enamel ornaments

worn

surface

worn

surface

horizontal

striations

worn

surface

unworn

enamel

unworn **

enamel m

worn

surface

worn

surface

unworn

enamel

carina

tooth surface

3 mm

Figure 2. Wear patterns on representative teeth of NMV P252567, suggesting suction feeding in an aetiocetid. A, left upper canine or first

premolar; B, double-rooted postcanine 1; C, ?lower double-rooted postcanine 2; D, ?lower double-rooted postcanine 3; E, micro-computed

tomography cross section of postcanine 1, showing the depth and rounded edges of the horizontal striations (marked by large black arrows).

A-C are shown in lingual, labial and anterior/posterior view, D in lingual view only.

(piston-like) movements of the tongue and/or flows of water

laden with prey and sediment. The horizontal orientation of

these forces is consistent with some form of (presumably

benthic) suction feeding, as in Mammalodon and Odobenus,

with the presence of lingual abrasion as far anteriorly as the

incisors suggesting that suction was used for prey capture.

This interpretation holds irrespective of the age of the

individual, as mature ontogeny would have exaggerated enamel

wear without producing a heavy lingual bias or, particularly,

the characteristic deep horizontal striations. Nevertheless, the

intact enamel on the left upper incisor demonstrates that at least

some of the anteriormost teeth were protected from wear, e.g.

by being largely covered by gum tissue or by being located far

away from the main flow of prey and water. Baleen and tooth-

assisted filter feeding can almost certainly be excluded, given

that (i) baleen was most likely absent (see below) and (ii) the

highly worn teeth would have been exceedingly poor at

retaining small food particles. There is also no clear evidence

for raptorial feeding, such as pronounced apical wear or

dorsoventral shear facets, although such features may have

been obliterated by heavy abrasion. At least facultative raptorial

feeding may therefore have been possible.

We are not aware of a modern marine mammal showing a

pattern of labial ‘hourglass’ wear that resembles that of NMV

P252567. Nevertheless, the anterior, posterior and labial wear

of the individual teeth is consistent with water and abrasive

Suction feeding preceded filtering in baleen whale evolution

Figure 3. Additional teeth of NMV P252567. A, left upper incisor; B, right upper incisor; C, double-rooted postcanine 4. All teeth are shown in

lingual (left) and labial (right) views. The lack or comparatively small degree of wear on the incisors suggests they may have been largely (left

upper incisor) or partially (right upper incisor) enclosed within the gingiva, protecting them from the abrasive wear that affected the other teeth.

A fifth double-rooted postcanine closely resembles postcanines 2 and 4 in terms of its wear, but is still partially encased in sediment and hence

not shown here.

particles being forcibly expelled from the oral cavity through

the diastemata. Similar water expulsion behaviour following

suction has been observed in living species, such as pilot

whales, belugas, leopard seals and Australian fur seals

(Hocking et ah, 2013; Hocking et al., 2014; Kane and Marshall,

2009; Werth, 2000a). During water expulsion, the jaws would

likely have been held slightly open, causing the nearly

occluding, interdigitating tooth rows to form a series of small

gaps defined by the rims of the individual diastemata and the

tips of the occluding upper or lower teeth. Sediment-laden

water forced through these gaps would have abraded the

enamel both along the rim of each diastema and on the

immediately adjacent, labial portions of the tooth crowns.

Over time, the labial wear surfaces would have enlarged into

the hourglass wear observed here, possibly aided by the

accidental, temporary retention of some sediment particles

inside the lips after each water expulsion event.

Did aetiocetids have baleen?

Aetiocetids have previously been proposed as the most basal

mysticetes to possess baleen, the key adaptation of modern

whales. More specifically, the widespread occurrence of

palatal nutrient foramina (in Aetiocetus, Fucaia and

Morawanocetus ), which in extant mysticetes supply the baleen

rack, has been used to infer the existence of an incipient baleen

structure between or just lingual to the teeth (Demere and

Berta, 2008; Demere et al., 2008). While such an interpretation

is possible, it also remains untested: just as the origin of

feathers in non-avian dinosaurs does not mark the beginnings

of flight, so the appearance of palatal foramina in mysticetes

need not indicate the presence of baleen. Instead, the foramina

of aetiocetids could, for example, have supplied its immediate

predecessor - namely, well-developed gums, the presence of

which is indicated both by the strongly emergent teeth of early

mysticetes (Demere and Berta, 2008; Fitzgerald, 2010) and,

possibly, the largely unworn incisor of NMV P252567.

The presence of palatal foramina in NMV P252567 cannot

be determined owing to post-mortem breakage of the rostral

margin. Nevertheless, this specimen is the first aetiocetid

preserving clear evidence of its feeding strategy, and thus also

the first test of the idea that baleen occurred in members of

this family. In the case of NMV P252567, extreme lingual

wear indicates that the teeth were directly exposed to strong

abrasive forces uninhibited by adjacent keratinous tubules or

plates. The deep horizontal striations in particular suggest that

the teeth were affected by continuous, linear movements of the

tongue and/or prey-laden water, which would have been

hindered if baleen had shielded the inside of the tooth row.

F.G. Marx, DP. Hocking, T. Park, T. Ziegler, A.R. Evans & E.M.G. Fitzgerald

The presence of baleen is made even less likely by the

interdigitating dentition, as judged from the mandibular

alveoli alternating with similarly-sized embrasure pits for

the upper teeth (fig. 1). Interdigitating teeth also occur in

Fucaia goedertorum (Barnes et al., 1995) and Aetiocetus

weltoni (Demere and Berta, 2008), and suggest a lack of

space and risk of functional interference (i.e. teeth potentially

damaging or disorganising the baleen rack) that speaks

against the presence of functional baleen. Overall, we

therefore conclude that NMV P252567 did not possess baleen

and was hence incapable of filter feeding in a manner similar

to modern mysticetes.

The condition of NMV P252567 reinforces previous, less

decisive evidence against baleen in several other aetiocetids,

such as the well-developed shear facets on the teeth of Fucaia

buelli and the large size of the teeth in both F. buelli and

Morawanocetus (Marx et al., 2015; Sawamura, 2008;

Sawamura et al., 2006). Specifically, shearing in F. buelli

would likely have posed a considerable risk of damage to the

baleen rack after each bite, while the relatively elongate teeth

of Morawanocetus (and, probably, F. buelli) result in short

diastemata, abrogating the need for a baleen filter. Both of

these observations rely on indirect evidence, but the difficulties

in explaining how baleen could have functioned in these taxa

are suggestive.

In extant baleen whales, tall lower lips, marked lateral

bowing and longitudinal (alpha) rotation allow the mandibles

to occlude on to the labial (rather than the ventral) surface of

the baleen plates, thereby preserving the integrity of the rack

(Lillie, 1915) (fig. 4). In Aetiocetus and Fucaia , essentially

straight mandibles, a tall, straight coronoid process, embrasure

pits, and the presence of attritional wear on the teeth (Demere

and Berta, 2008; Emlong, 1966) demonstrate that the lower

jaw moved largely vertically and was positioned close to the

upper jaw to enable tooth occlusion (fig. 4). An aetiocetid

baleen rack would have been closely associated with the teeth,

as judged from the position of the palatal foramina in A.

weltoni and the juxtaposition of the rudimentary teeth and

developing baleen in extant mysticete foetuses (Demere et al.,

2008; Ishikawa and Amasaki, 1995). As a result, aetiocetid

baleen would have been constantly disturbed by mandibular

contact.

Teeth could conceivably have acted as protective spacers

between the jaws, allowing baleen to grow between or just

medial to the upper teeth. However, the interdigitating

dentition would still have resulted in considerable disturbance

of the rack. It is also possible that the inherent flexibility of

baleen would have allowed it to withstand compression, e.g. by

folding away posteriorly as in extant bowhead whales (Werth,

2001; Werth, 2004). Unlike in bowhead whales, however, the

presence of teeth in aetiocetids - both adjacent to the rack and

coming from below - would likely have interfered with the

folding process. We therefore suggest that, contrary to past

proposals (Demere and Berta, 2008; Demere et al., 2008), the

evolution of baleen likely only became feasible after the

appearance of a laterally bowed mandible capable of clearing

the baleen rack during mouth closure, and likely following the

reduction or loss of emergent dentition (fig. 5).

Current model of baleen evolution

Current ideas on the origin of baleen argue for a direct

transition from raptorial to bulk filter feeding, as seemingly

exemplified by aetiocetids in their retention of functional teeth

alongside features generally associated with filtering (Demere

et al., 2008). Besides the presence of (i) palatal foramina, these

features include (ii) thin lateral margins of the maxillae; (iii) a

relatively broad rostrum; and (iv) an unsutured, ligamentous

mandibular symphysis. Laterally bowed mandibles, another

feature claimed to be present in aetiocetids (Demere et al.,

2008), is not apparent in any of the specimens we examined

(NMV P252567, fig. IB; Aetiocetus weltoni, UCMP 122900;

Fucaia goedertorum, LACM 131146), all of which instead

possess effectively straight lower jaws.

While it is true that these traits facilitate bulk filter feeding

in modern mysticetes (e.g. Lambertsen et al., 1995), their

condition and function in archaic mysticetes is much less

clear. As argued above, evidence from NMV P252567 and

other aetiocetids speaks against the presence of baleen in this

family, with the palatal foramina - the prime evidence for

baleen - more likely supplying enlarged gums. Likewise, we

see no direct link between thin lateral maxillary margins and

filtering, and instead suggest that they may be a consequence

of rostral broadening. The resulting increase in oral capacity

would benefit both suction performance and filter feeding, so

cannot be attributed to filtering alone (Fitzgerald, 2012; Werth,

2006). In any case, broad rostra are not characteristic of all

filter-feeding whales: those of skim-feeding right whales are

narrow and elongate, as essentially are those of the extant

pygmy right ( Caperea marginata ) and even grey whales

(Eschrichtius robustus).

Finally, the exact dental occlusion and tall, straight coronoid

processes of Fucaia and Aetiocetus imply that longitudinal

(alpha) rotation of aetiocetid mandibles was minimal compared

with extant mysticetes, despite a ligamentous symphysis (Kimura,

2002; Lambertsen et al., 1995; Marx et al., 2015). Among extant

mysticetes, a ligamentous mandibular symphysis enables extant

balaenopterids to rotate their bowed mandible along its long axis,

thereby increasing oral volume during engulfment feeding

(Lambertsen et al., 1995). By contrast, the mandible of aetiocetids

is straight and constrained to largely dorsoventral rotation,

rendering a mobile symphysis ineffective for increasing oral

capacity (Arnold et al., 2005; Marx et al., 2015).

Alternatively, mandibular rotation may initially have

enhanced control of the lower lip. In right whales, lateral lip

rotation serves to create a flow channel lateral to the baleen

rack during skim feeding (Lambertsen et al., 2005; Werth and

Potvin, 2016). This feeding strategy requires a large filtration

area, which in right whales is created by the arched rostrum

and elongate baleen plates. Given its short, flat rostrum and

erupted teeth, space limitations in the aetiocetid skull would

have precluded this feeding mode. In grey whales, lip rotation

appears to assist lateral suction feeding by creating an aperture

for prey and water to be sucked into the oral cavity (Ray and

Schevill, 1974). A similar behaviour in aetiocetids is

conceivable, but the tall coronoid process would likely have

prevented the opening of a wide enough gap.

Suction feeding preceded filtering in baleen whale evolution

Figure 4. Cross section of the rostrum and lower jaws of A, a balaenid, B, a balaenopterid, and C, an aetiocetid, illustrating the relative movement of

the mandible during jaw closure (red arrows). All drawings show the mouth slightly open. In right whales (A) and rorquals (B), the laterally bowed

mandibles and/or tall lower lips rotate inwards on to the labial surface of the baleen plates, thereby leaving the rack intact. In aetiocetids (C), the

movement of the mandible is mostly vertical and the upper and lower jaws need to approach each other enough to allow the teeth to occlude, thereby

risking interference with any baleen present. A and B are adapted from Pivorunas (1979: fig. 3).

"^=i enlarged gingiva (palatal nutrient foramina)

I cga laterally bowed mandibles

i baleen

* i complete loss

t of teeth

Eomysticetidae

crown Mysticeti

Basilosauridae

Aetiocetidae

suction + simple sieving

suction + baleen filtering

ram + baleen filtering

, Mysticeti_

| Chaeomysticeti (teeth reduced or absent, baleen-bearing)

Figure 5. Suction feeding precedes baleen filtering in mysticete evolution. A, consensus tree of aetiocetid evolutionary relationships, based on all

cladistic studies published to date(e.g. Demere and Berta, 2008; Demere et al., 2008; Fitzgerald, 2010; Geisler and Sanders, 2003; Marx and Fordyce,

2015; Steeman, 2007), showing major feeding-related synapomorphies; B, life reconstructions (top) and skulls (in lateral view) of a representative

archaeocete (Dorudon atrox), aetiocetid (NMV P252567), eomysticetid (Yamatocetus canaliculatus) and extant suction feeding mysticete (grey

whale, Eschrichtius robustus)-, C, inferred behaviours and feeding strategies. Life reconstructions by Carl Buell.

F.G. Marx, DP. Hocking, T. Park, T. Ziegler, A.R. Evans & E.M.G. Fitzgerald

Overall, the function of the ligamentous symphysis in

aetiocetids remains unclear. In particular, there is little

evidence to suggest it was a specific adaptation to filter feeding.

This point is emphasised further by the observations that, even

among extant mysticetes, jaw rotation can be associated with

suction feeding rather than filtration, and that an unsutured

symphysis also occurs in a variety of other mammals,

including - possibly - Mammalodon (Fitzgerald, 2010;

Lieberman and Crompton, 2000).

An alternative model of baleen evolution

NMV P252567 makes a crucial contribution to the question of

how baleen and bulk filtering first evolved. It is one of a limited

number of fossil whales documenting the transition from

raptorial to filter feeding; its cranial morphology disputes

prior conjecture about the widespread presence of baleen in

aetiocetids; and it provides the first reported evidence of

suction feeding at the pivotal mysticete transition towards

filter feeding and giant size. The ability to generate suction is

fundamental to most marine vertebrates, and widespread

among extant marine mammals, including pinnipeds and

cetaceans (Hocking et al., 2013; Hocking et al., 2014; Kane

and Marshall, 2009; Werth, 2000b; Werth, 2006). Nevertheless,

up to this point it has rarely been associated with mysticete

evolution, other than in reference to the highly unusual

mammalodontids (Fitzgerald, 2010; Fitzgerald, 2012).

Suction is necessary when feeding underwater, where it

enables the transport of food towards the back of the mouth for

swallowing even in raptorial species that still employ teeth in

prey capture (Werth, 2000b; Werth, 2006). This was likely

already the case in archaic whales, soon after their initial

transition to an aquatic environment. However, suction

behaviour - whether for prey capture or intraoral transport - is

generally difficult to demonstrate in fossils, since relevant

osteological correlates, such as blunt, wide jaws (Werth, 2006)

or a large hyoid apparatus (Bloodworth and Marshall, 2007;

Heyning and Mead, 1996), are often either not preserved, or

not always clearly developed, such as in the grey whale,

Eschrichtius robustus (Kienle et al., 2015).

NMV P252567 offers an extremely rare insight into the

evolution of suction behaviour and, along with Mammalodon,

demonstrates a tendency for early mysticetes to evolve suction-

based feeding strategies. There is currently no evidence that

other aetiocetids relied on suction to a similar degree, although

such a behaviour may be less apparent in animals that feed

higher in the water column, and hence ingest less or no

abrasive (i.e. wear-inducing) sediment. Nevertheless, given the

apparently high degree of specialisation of NMV P252567 and

the widespread occurrence of suction behaviour among extant

marine mammals, it seems highly likely that aetiocetids were

at least able to use suction for intraoral transport.

Use of suction and lack of baleen in aetiocetids suggests an

alternative model - briefly hinted at by Arnold et al. (2005) - of

how and why filter feeding first arose (fig. 5). Archaic mysticetes,

including aetiocetids, likely inherited both a functional dentition

and the ability to use suction for intraoral transport from their

archaeocete ancestors (Werth, 2000b). Water ingested as a

result of suction was expelled prior to swallowing (Hocking et

al., 2013; Hocking et al., 2014; Kane and Marshall, 2009; Werth,

2000a), with the prey either being physically held in place, or

the teeth, jaws and surrounding soft tissues acting as a barrier,

or simple sieve, retaining food items inside the mouth

(Bloodworth and Marshall, 2005; Hocking et al., 2013). Some

of these early whales, including NMV P252567, Mammalodon

and the ancestor of modern mysticetes, honed their suction

capabilities to the point where they became able to capture prey,

and we suggest that it was this transition, not filter feeding, that

ultimately initiated tooth loss in the chaeomysticete lineage.

Among both extant (sperm whales, beaked whales and

certain delphinids) and extinct odontocetes (e.g.

Australodelphis, Odobenocetops ), capture suction feeding

strongly correlates with a reduced dentition (Werth, 2000b;

Werth, 2006), and the same may plausibly have been the case

in mysticetes. This scenario avoids potential problems of

functional interference between a working dentition and

incipient baleen (Marx et al., 2015), and explains how teeth

could have been lost without impacting on foraging success.

Further, a loss of functional teeth prior to the origin of baleen

coincides with evidence of foetal development from extant

mysticetes, which shows that baleen growth only initiates once

the tooth buds have already started to degrade (Ishikawa and

Amasaki, 1995; Karlsen, 1962). It is possible that teeth and

baleen nonetheless co-occurred in some archaic

chaeomysticetes, as shown by eomysticetids bearing shallow

alveoli and, possibly, teeth (Boessenecker and Fordyce, 2015);

however, the dentition in these taxa was already reduced. We

also note the similar anterior positioning of teeth in

eomysticetids and extant suction-feeding odontocetes like the

beluga, and the delphinids Grampus and Globicephala.

Suction for capture limited the maximum size of prey that

could be taken, and furthermore would have enabled the

ancestors of modern mysticetes to gather small prey items in

bulk; however, the absence of specialised filtering teeth, such

as those of the extant crabeater ( Lobodon ) and leopard seals

(Hydrurga ), would have permitted the inadvertent expulsion of

small food particles prior to swallowing, as observed in trials

with California sea lions ( Zalophus californianus ) (Hocking et

al., 2013). This problem was eventually solved by the elaboration

of the gingiva, first potentially as a grasping (Miller, 1929) and,

ultimately, a filtering apparatus - i.e. baleen. A similar

condition exists in the extant Dali’s porpoise Phocoenoides

dalli, which supplements its rudimentary dentition with a series

of ‘gum teeth’ that are structurally similar to the early growth

stages of baleen (Miller, 1929). As Miller (1929: 4) himself

observed: “These resemblances are so important that we are

probably justified in regarding the gingival and dental structures

of Phocoenoides as representing anatomical stages closely

parallel to those through which the corresponding parts in the

toothed ancestors of the Mysticeti must have passed.”

The feeding strategy of the earliest baleen-bearing whales

would initially have been a form of intermittent or continuous

suction filter feeding, as inferred for a range of extinct

cetotheriids (El Adli et al., 2014; Gol’din et al., 2014), and still

observed in the extant grey whale, Eschrichtius robustus (Ray

and Schevill, 1974). However, with baleen now in place, other

methods of filtering no longer reliant on suction also became

Suction feeding preceded filtering in baleen whale evolution

possible, including the highly specialised skim (Werth and

Potvin, 2016) and lunge feeding (Lambertsen et al., 1995)

strategies of extant right whales and rorquals, respectively.

Our new model is consistent with all available

palaeontological, developmental and behavioural evidence,

but will benefit from further research effort. This might

include an investigation of dietary stable isotopes, to determine

at what trophic level aetiocetids were feeding (e.g. Clementz et

al., 2014); an increased focus on the oldest (Late Eocene-Early

Oligocene) mysticetes, to test for evidence of suction feeding

in early chaeomysticetes (e.g. tooth wear), or further evidence

regarding baleen in aetiocetids, e.g. in the form of actually

preserved traces (e.g. Esperante et al., 2008; Gioncada et al.,

2016); and further studies of the feeding strategies of extant

marine mammals, to determine possible modern analogues of

archaic mysticetes. Overall, our findings suggest that suction

behaviour was fundamental to the evolution of baleen and

filtering, and thus a crucial early innovation that helped to

trigger the rise of the largest animals on Earth.

Acknowledgements

We thank J.L. Goedert, S.R. Benham and D. Reed for

collecting and donating the specimen, B. Francischelli, D.

Pickering, J. Rule and A. Werner for their critical contributions

to its preparation, A. Collareta, T. Kimura and O. Lambert for

their constructive reviews of our manuscript, and Carl Buell

for providing life reconstructions of a range of extinct and

extant cetaceans. This research was supported by a Marie

Sklodowska-Curie Global Postdoctoral fellowship (656010/

MYSTICETI) to F.G.M., Australian Research Council Future

Fellowship FT130100968 to A.R.E., Australian Research

Council Linkage Project LP150100403 to A.R.E. and

E.M.G.F., and an Australian Postgraduate Award to T.P

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