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Two commercially valuable holothurians, the sandfish and golden sandfish, vary in colour and have a confused
taxonomy, lending uncertainty to species identifications. A recent molecular study showed that the putative variety
Holothuria (Metriatyla) scabra var. versicolor Conand, 1986 (golden sandfish) is a distinct species from, but could
hybridize with, H. (Metriatyla) scabra Jaeger, 1833 (sandfish). Examination of the skeletal elements and external
morphology of these species corroborates these findings. The identity of H. (M.) scabra is unambiguously defined
through the erection and description of a neotype, and several synonyms have been critically re-examined. The
nomenclaturally rejected taxon H. (Metriatyla) timama Lesson, 1830 and H. (M.) scabra var. versicolor (a nomen
nudum) are herein recognized as conspecific and are allocated to a new species, Holothuria lessoni sp. nov., for
which type specimens are described. The holotype and only known specimen of H. aculeata Semper, 1867, has been
found and is redescribed. It is considered to be a valid species. Taxonomic clarification of this heavily exploited
species complex should aid its conservation and permit species-specific management of their fisheries. 2009
The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059.
ADDITIONAL KEYWORDS: conservation biology Holothuria aculeata Holothuria lessoni sp. nov.
Holothuria scabra Holothuria scabra versicolor Holothuria timama nomenclature.
INTRODUCTION
Some 20 large sea cucumber species in the order
Aspidochirotida are fished commercially throughout
the Indo-Pacific. We suspect, however, that this is an
underestimate of the actual number of aspidochirotid
species exploited because several species are poorly
defined taxonomically. This absence of sound taxo-
*Corresponding author.
E-mail: [email protected]
Current address: Beechcroft, Norwich Road, Scole, Norfolk
IP21 4DY, UK.
40
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
41
RESULTS
SYSTEMATIC
ACCOUNTS
NOV.
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
42
C. MASSIN ET AL.
Figure 1. AD, Holothuria (Metriatyla) lessoni sp. nov. A, holotype (the covering fine layer of sand was gently
brushed away); B, holotype (top) and paratype (bottom); C, black form (IG 30768/5); D, mottled form (IG 30768/4). E, H.
timama Lesson, 1830, original drawing; F, H. timama Lesson, 1830, remaining fragment of holotype. Photographs A &
B by C. Massin, C & D by S. Purcell; E & F by Y. Samyn.
MATERIAL
EXAMINED
Type material
Holotype H. lessoni sp. nov.: RBINS IG 27754/179,
Hansa Bay (Madang Province, Papua New Guinea),
muddy bottom with sparse seagrass bed, 6-m depth,
coll. C. Massin 01.x.1990.
Paratype H. lessoni sp. nov: RBINS IG 27754/180,
same data as holotype.
Holotype H. timama Lesson, 1830 (buccal apparatus
only): MNHN EcHh 544, Waigeo Island, substrate
and depth not given, coll. unknown.
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
43
Figure 2. A, Holothuria (Metriatyla) aculeata Semper, 1868, holotype; B, H. aculeata, original drawing; C. Holothuria
(Metriatyla) scabra, non-type material, New Caledonia (IG 30768/1); D, Holothuria cadelli Bell, 1887, lectotype; E,
Holothuria gallensis Pearson, 1903, original drawing; E, Holothuria cf. gallensis Pearson, 1903, drawing by Selenka, 1867
as H. tigris Brandt, 1835; G, Holothuria saecularis Bell, 1887, paralectotype. Photograph A by A. Martynov; B, DG by
Y. Samyn; C. by S. Purcell.
Non-type material
RBINS IG 27598/133 (1 specimen): Hansa Bay
(Madang Province, Papua New Guinea), muddy
bottom with sparse seagrass bed, 11 m depth, coll. C.
Massin 13.x.1989.
RBINS IG 27598/135 (1 specimen): Hansa Bay
(Madang Province, Papua New Guinea), muddy
bottom with sparse seagrass bed, 10 m depth, coll. C.
Massin 13.x.1989.
RBINS IG 30768 (6 specimens; labelled RBINS IG
30768/2 to 30768/7): lot Matre (New Caledonia),
sandy bottom with sparse seagrass bed, 0.52 m
depth, coll. S. Purcell 23.ii.2007.
MAGNTD
Q004089: Ashmore
Reef
Lagoon,
121520S, 1225930E, stn. 935 (Australia), 1 m
depth, coll. L. Vail 10.iv.1987.
TYPE
LOCALITY
ETYMOLOGY
The name lessoni is given in honour of Dr R. P. Lesson
who, in 1830, was the first to recognize and describe
the species under the name Holothuria timama
(Fig. 1E). However, as the name timama, as published
in the binomen Holothuria timama, is suppressed
(ICZN Opinion 762, 1966), it can no longer be used to
denominate this species.
KNOWN
GEOGRAPHICAL DISTRIBUTION
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
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C. MASSIN ET AL.
Figure 3. Holothuria (Metriatyla) lessoni sp. nov., holotype. A, calcareous ring (R, radial plate; IR, interradial plate);
B, buttons of dorsal body wall; C, tables of dorsal body wall; D, large table of dorsal body wall; E, tables of ventral tube
feet; F, buttons of ventral tube feet; G, perforated plates of ventral tube feet. Scale bars: A = 5 mm; BG = 50 mm.
TAXONOMIC
DESCRIPTION
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
45
Figure 4. Holothuria (Metriatyla) lessoni sp. nov., holotype. A, perforated rods of ventral tube feet; B, perforated
plates of ventral tube feet; C, perforated plates of dorsal tube feet; D, perforated plates and rods of dorsal tube feet; E,
tables of dorsal papillae; F, reduced end plate of a dorsal papilla; G, perforated plates of the anal papillae; H, perforated
plates of cloacal wall. Scale bars: A, D & G = 100 mm; B, C, E, F & H = 50 mm.
occasionally have black patches ventrally. Undisturbed animals from Papua New Guinea covered dorsally by thin layer of sand. Body wall smooth to the
touch, 1015 mm thick in contracted specimens. Calcareous ring very stout (Fig. 3A), radials twice as
wide as inter-radials, radials with deep anterior
notch. Tentacle ampullae 2050 mm long in preserved
material. One Polian vesicle, 3037 mm long in preserved material. Stone canal single, very long, 150
190 mm long in preserved material. Gonad well
developed, with numerous, very long, non-branching
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
46
C. MASSIN ET AL.
Figure 5. Holothuria (Metriatyla) lessoni sp. nov., holotype. A, rods and small perforated rods of the cloacal wall; B,
tentacle rods; C, detail of the extremities of large tentacle rods. Scale bars: A = 50 mm; B = 200 mm; C = 100 mm.
Table 1. Distribution of the golden sandfish, H. (M.)
lessoni sp. nov., based on scientific literature and credible reports
Locality
Reference
New Caledonia
Papua New Guinea
Torres Strait
Great Barrier Reef
Moreton Bay
Western Australia
Indonesia
East Indies
Seychelles
Madagascar
East Africa (Kenya)
REMARKS
It is usually possible to distinguish between the
species lessoni and scabra in the field (no field data
for H. aculeata are available) based on colour patterns, form and length of papillae/tube-feet and
presence/absence of wrinkles in the body wall (see
Table 2). As these characters are known to show some
variation (see below), the identification of each species
must rely on the study of the ossicle assemblage. The
most striking differences are: (1) the edge of the disc
of the tables in H. lessoni has a spiny rim, whereas H.
aculeata and H. scabra have tables with a smooth
rim, and (2) the cloacal wall of H. lessoni has irregular rods, while these are lacking in H. aculeata and
H. scabra.
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
MATERIAL
EXAMINED
Type material
Holotype (labelled Unicum on the original label):
ZMMSU H115.
47
Non-type material
None.
TYPE
LOCALITY
Bohol, Philippines.
KNOWN
GEOGRAPHICAL DISTRIBUTION
TAXONOMIC DESCRIPTION
Gross morphology: Medium-sized holothurian; preserved specimen 95 mm long and 35 mm across,
cylindrical, with transverse ridges; dimensions in
life 22 cm long and 6 cm wide at mid body (according to Semper, 1868: pl. 24). Mouth ventral, surrounded by (at least) 17 brown tentacles and a
collar of well-developed papillae. Anus terminal,
anal papillae not observed. Colour in alcohol: dorsal
surface cream with irregular brown dots, ventral
surface whitish, tube feet deep brown (Fig. 2A).
Colour in life (according to Semper, 1868): ventral
surface uniform whitish; dorsal surface darker,
somewhat reddish with irregular dark brown
patches; tube feet darker than background colour
(Fig. 2B). Dorsal appendages not aligned in rows.
Ventral tube feet sparse, spread evenly over trivium.
Body wall gritty to the touch, 49 mm thick. Original dissection of holotype makes observation of calcareous ring detrimental to unique specimen.
Tentacle ampullae short, 710 mm long. Polian
vesicles, two, very large (19 and 20 mm long). Stone
canal, single, large, 12 mm long. Gonad branched.
Longitudinal muscles very large, thick, lateral sides
not attached to body wall, margins heavily curled
upwards, giving muscles U-shaped form. Respiratory
trees present. Cuvierian tubules absent. Digestive
tract missing except a small part of anterior dorsal
lobe.
Ossicles: Dorsal body wall with tables and buttons:
tables with disc quadrangular, smooth, perforated by
four central holes and one circle of peripheral holes,
pillar short, united by one cross-beam ending in
crown of short blunt spines (Fig. 6A), majority of
tables 50 mm across and 30 mm high, few large ones
with two circles of peripheral holes and up to 80 mm
across (Fig. 6B); buttons with 34 pairs of holes,
very knobbed, 4060 mm long (Fig. 6C). Dorsal tube
feet with tables, few buttons, perforated plates and
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
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C. MASSIN ET AL.
Figure 6. Holothuria aculeata Semper, 1868, holotype. A, tables of dorsal body wall; B, large table of dorsal body wall;
C, buttons of dorsal body wall; D, table of dorsal papillae; E, perforated plates of dorsal papillae; F, rods of the dorsal
papillae; G, buttons of dorsal papillae; H, table of ventral body wall; J, buttons of ventral body wall; K, table of ventral
tube foot; L, buttons of ventral tube feet; M, perforated plates and rods of ventral tube feet. Scale bars: AM = 50 mm.
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
49
Figure 7. Holothuria aculeata Semper, 1868 holotype. A & B, tentacle rods; C, ossicles of cloacal wall. Scale bars:
A = 100 mm; B & C = 50 mm.
REMARKS
Despite several records in the literature (cf. synonymy list above), either as a valid species, as a
misidentified species or as a synonym of H. timama
Lesson, 1830 (= H. lessoni sp. nov.), virtually no information directly attributable to this form subsequent
to the original description of H. aculeata exists. This
is due, at least in part, to the fact that the type
material of H. aculeata was considered lost (cf. Rowe
& Gates, 1995) as it could not be located in any of the
German museums that hold type material of Semper
(C. Lter, pers. comm.). However, through Drs A.
Smirnov and A. Martynov, we were able to locate the
material in the ZMMSU. To our knowledge, this specimen the holotype as the label bears the denomination Unicum is the only available voucher. This
implies that our knowledge of the intraspecific variety
of this species is non-existent.
The holotype presents irregular dorsal black areas
as do some specimens of H. lessoni sp. nov. In spite of
this, the body wall ossicles of H. aculeata are closer to
those of H. scabra than to those of H. lessoni.
However, H. aculeata differs from H. scabra by the
colour pattern (dorsal surface with dark irregular
patches versus uniform grey green colour, respectively), by the large tables of the dorsal body wall (up
to 80 mm across with normal pillars versus up to
120 mm across with reduced pillars, respectively) and
by the ossicles of the cloacal wall (4065 mm across
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
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C. MASSIN ET AL.
Figure 8. Holothuria scabra Jaeger, 1833, neotype. A, calcareous ring (R, radial plate; IR, interradial plate); B, buttons
of ventral body wall; C, tables of ventral body wall; D, tables of dorsal body wall; E, buttons of dorsal body wall; F, large
table of dorsal body wall; G, buttons of ventral tube feet; H, tables of ventral tube feet; J, perforated rods of ventral tube
feet; K, end plate of ventral tube foot; L, perforated plates of ventral tube feet. Scale bars: A = 5 mm; BH, L = 50 mm; J
& K = 100 mm.
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
51
Figure 9. Holothuria scabra Jaeger, 1833, neotype. A, tables of dorsal papillae; B, buttons of dorsal papillae; C,
perforated rods of dorsal papillae; D, perforated rods of anal papillae. Scale bars: AD = 50 mm.
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
52
C. MASSIN ET AL.
MATERIAL
EXAMINED
Type material
Neotype of H. scabra (designated here): RBINS IG
28251/36, Panikiang (Sulawesi, Indonesia), reef flat
at low tide, coll. C. Massin 30.viii.1994.
Four syntypes of Holothuria cadelli Bell, 1887:
NHM 1886.6.26.8692, Andaman Islands, coll. Dr
Anderson, date unknown; the largest specimen
(180 mm long) is here designated as lectotype.
Two syntypes of Holothuria saecularis Bell, 1887:
NHM 1866.4.13.5, Angola, coll Dr Welxitch, date
unknown. the largest specimen (180 mm long) is here
designated as lectotype.
Non type material
RMNH Ech 6081 (one specimen): Panikiang
(Sulawesi, Indonesia), reef flat at low tide; coll. C.
Massin 30.viii.1994.
RBINS IG 28767 (ten specimens from aquaculture):
Vonavona lagoon (Kohinggo Island, Western Province,
Solomon Islands), 2-m depth, coll: J. F. Hamel
ii.1998.
RBINS IG 30768/1 (one specimen): lot Matre
(New Caledonia), on a sandy bottom with sparse
seagrass beds at 1-m depth; coll: S. Purcell 27.ii.2007.
TYPE
LOCALITY
KNOWN
GEOGRAPHICAL DISTRIBUTION
TAXONOMIC
DESCRIPTION OF NEOTYPE
Gross morphology: Medium-sized holothurian; preserved neotype 175 mm long and 40 mm wide. Body
arched dorsally and more or less flat ventrally; one
median longitudinal groove ventrally. Mouth ventral,
surrounded by 20 short tentacles. Anus terminal with
anal papillae. Dorsal surface grey-green with transverse greenish bands (Fig. 2C). Ventral surface greywhite, speckled with dark tiny dots corresponding to
tube feet. Body wall gritty to the touch, thin (23 mm
thick). Calcareous ring with radial plates twice as
wide as interradial pieces. Radial pieces with deep
anterior notch (Fig. 8A). Stone canal single, 20 mm
long. One Polian vesicle, 32 mm long. Tentacle ampullae nearly as long as Polian vesicle. Longitudinal
muscles thin, flat with free extremities. No Cuvierian
tubules. Gonads not observed. Respiratory trees
simple, reaching up to calcareous ring. Digestive tract
with a long loop, filled with coarse sand.
Ossicles: Dorsal and ventral body wall with buttons
and tables. Ventrally buttons very numerous, comparatively large, 4075 mm long (Fig. 8B); tables rare,
4055 mm high with disc 6095 mm across, quadrangular, smooth, perforated by one central hole and one
circle of 816 peripheral holes; spire of four short
pillars united by one cross-beam, and ending in a
crown of blunt spines (Fig. 8C); crown never as wide
as table disc. Dorsally, tables very similar to ventral
ones (Fig. 8D); buttons nodulous, 4050 mm long, with
34 pairs of holes (Fig. 8E); a few large buttons with
57 pairs of holes (Fig. 8E). Dorsally, a few large
tables (Fig. 8F), 8090 mm across (up to 120 mm across
in other specimens) with numerous holes and with
reduced pillars. Ventral tube feet with nodulous
buttons, 4090 mm long (Fig. 8G); perforated rods,
110175 mm long (Fig. 8J); and tables, 50100 mm
across (Fig. 8H), very similar to those of body wall;
end plate 400460 mm across with a second layer
(Fig. 8K), and surrounded by spiny perforated plates
(Fig. 8L). Dorsal papillae with few rods (Fig. 9C) and
numerous buttons nearly all with three pairs of holes
and similar to those of tube feet (Fig. 9B); tables rare
(Fig. 9A) or absent; end plate more or less 270 mm
across, comprising several pieces. Anal papillae with
tables, buttons and rods; rods slightly curved, 130
200 mm long with few perforations at extremities and
sometimes 12 central perforations (Fig. 9D). Tables
60100 mm across with a quadrangular, spiny disc
and often an irregular crown of spines. Cloacal wall
with large spiny perforated plates 80140 mm across
with 417 holes (Fig. 10A); most of the plates with
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
REMARKS
The neotype of H. scabra has a particularly thin body
wall in comparison with larger specimens of the
species. This body wall thickness is in agreement with
the lengthweight relationship of H. scabra. Specimens grow generally to 150170 mm in length; larger
specimens become heavier by an increase of body-wall
thickness (Pitt & Nguyen, 2004). Colour in live specimens can be quite variable. Entirely black morphs
have been observed on Australias east coast; these
were genetically indistinguishable from the green to
grey morphs (Uthicke & Benzie, 1999). We also
observed specimens with chocolate brown dorsal sides
from Bali (Indonesia; S. Uthicke unpubl. data). Black
morphs of H. scabra are also quite common in New
Caledonia (S. W. Purcell, pers. observ.).
The species Holothuria cadelli Bell, 1887, H. gallensis Pearson, 1903 and H. tigris Brandt, 1835 have
been included in the synonymy at least since Pannings (1935) report. Considering the confusion surrounding the true concept of the species scabra, as
demonstrated above, the type series of cadelli was
re-examined. Type specimens of both gallensis and
tigris have not been located, although the original
descriptions have served well in understanding their
external morphology. The type specimen of a fourth
species, H. saecularis Bell, 1887, listed by Panning as
a valid species (but poorly described) has also been
examined and we herein are able to commit it to the
synonymy of H. scabra.
To clarify the relationship between each of these
species and H. scabra we briefly discuss them, in
alphabetical order, below.
Holothuria cadelli Bell, 1887
Decision for synonymy: Panning (1935).
Type data: Four syntypes, NHM 1866.6.26.8692 with
type locality Andaman Islands; largest specimen
(180 mm long, NHM 1886.6.26.86a) here designated
as lectotype; paralectotypes with collection numbers
NHM 1886.6.26.86bd.
Taxonomic description (see also Bell, 1887a): Type
material well preserved, well relaxed. Largest specimen 180 mm long and 40 mm wide at mid body. Body
cylindrical with rather fusiform extremities, espe-
53
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
54
C. MASSIN ET AL.
1994: 26, fig. XI; Conand, 1999: 23, pl. 3F; Samyn,
2003: 149, pl. 3A). It must be noted that these illustrations all are of specimens from the Indian Ocean.
The taxonomic status of H. gallensis will remain
doubtful until more material from the Indian Ocean
becomes available for morphological and molecular
determination. If the Indian Ocean form does prove to
represent a valid species then the name gallensis is
the oldest subjective synonym available, but needs to
be stabilized, taxonomically, by the establishment and
description of a neotype.
Holothuria tigris Brandt, 1835
Taxonomic decision for synonymy: Panning (1935).
Type data: Status and whereabouts undetermined;
type locality Uleai Is., Caroline Archipelago.
Taxonomic description: Cf. Brandt, 1835: 55 (in
Latin).
Remarks: The original description clearly states that
the dorsum has transverse dorsal stripes that can be
interrupted. As this conforms with H. scabra, H. tigris
best remains a junior subjective synonym.
Holothuria saecularis Bell, 1887
Type data: Two syntypes, NHM 1866.4.13.5 with type
locality Angola; largest specimen (150 mm long, NHM
1866.4.13.5a) here designated lectotype; paralectotype with collection number NHM 1866.4.13.5b.
Taxonomic description: Cf. Bell, 1887b: 534, pl. XLV,
fig. 6.
Examination of the gross morphology and the
ossicle assemblage leaves no doubt that this name
should be put in the synonymy of H. scabra. The type
locality (Angola, West Africa) is, however, very problematic as this implies that H. scabra is also present
in the Atlantic Ocean. We can only conclude that this
is an error of Bell.
Remarks: Bell (1887b) stated that no tables were to be
found; our (re-)examination, however, did reveal
tables, in size and shape identical to those of H.
scabra.
DISCUSSION
It is remarkable that the taxonomy of one of the most
valuable of tropical commercial species has remained
so chaotic, despite recent monographs on the species
(Bai, 1980; Hamel et al., 2001). The main problem has
been the lack of name-bearing types for two of
the three Indo-Pacific sandfish species now recog-
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
55
H. lessoni
H. aculeata
H. scabra
Body length
Body shape
250480 mm
Distinctly flattened
ventrally
Beige-brown or black,
mottled or blotched,
lighter ventrally
No transverse body
wrinkles, or only on
average 0.5 mm
(0.3 SD) deep
Black, bordered by a black
circle
Numerous (30 cm-2)
95 mm
Mildly flattened ventrally
150200 mm
Mildly flattened ventrally
Blotched
Grey-brown, grey-green, or
black, with or without
dark transversal bands
Transverse body wrinkles,
on average 3.1 mm
(0.7 SD) deep
Not applicable
External coloration
Coloration of dorsal
appendages
Distribution of dorsal
appendages
Length dorsal appendages
(living specimens)
Table: disc edge
Table diameter
Large table with reduced
pillars
Table height
Table crown
Perforated plate of cloaca
Rods of cloaca
Anal papillae rod shape
5585 mm
Large, acute spines
60130 mm, 9 to > 30 holes
Present, 60130 mm
Numerous, with large
central perforated process
Not applicable
Deep brown
Sparse (15 cm-2)
Smooth
4555 mm
Up to 80 mm
4045 mm
Small, blunt spines
4065 mm, 24 large holes,
Absent
Anal papillae not observed
ACKNOWLEDGEMENTS
We thank the many people who have helped us with
collecting information and making type material
available. In particular we would like to express our
gratitude to Dr N. Cominardi of the MNHN for granting us access to the sandfish types deposited in her
museum, Dr A. Cabrinowic for providing access to
type material in the NHM, Dr C. Lter of the ZMB
and Dr A. Smirnov of the Zoological Institute of
Saint-Petersburg for helping us in the search for the
type of H. aculeata, Dr A. Martynov of the ZMMSU
for the loan of the holotype of H. aculeata, Dr D.
Lagunov of the Manchester Museum for carrying the
type specimen of H. aculeata from Russia to England,
and Drs A. Polaszek and S. Tracey of the ICZN for
clarifying a nomenclatural issue. The expedition to
Papua New Guinea was financially supported in 1989
by the Belgian Fund for Basic Research (grant no.
209001.86) and by the Royal Belgian Institute of
Natural Sciences, and in 1990 by the King Lopold III
2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059
56
C. MASSIN ET AL.
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2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 4059