Zoologica Scripta, Vol. 20, No. 3, pp. 301-306, 1991 0300-3256/91 $3.00 .

00 +
Printed in Gr eat Britain Pergamon Press plc
01991 The Norwegian Academy of Science a nd Letters

ln situ observations on the genus Bathyplotes

(Holothuroidea, Echinodermata) in Antarctica and its
relevance to taxonomy
JULIAN GUTT and MICHAEL KLAGES
Accepted 11 September 1990

Gutt, J. & Klages, M . 1991. In situ observations on the genus Bathyplotes (Holothuroidea,
Echinodermata) in Antarctica and its relevance to taxonomy.-Zool. Scr. 20: 301-306.

Nineteen holothurian specimens were photographed or videotaped in situ on the deeper shelf of
the Weddell Sea (Antarctica). These specimens show a different gradient and combinations of
characteristics which are found in Bathyplotes rubipunctatus Gutt, 1990. A close relationship to this
species is therefore suggested. However, based on external features which are conspicuous in live
animals but not visible in preserved material they are classified into five groups. Further
investigations will reveal whether these groups are distinct species, or merely variants of B.
rubipunctatus.

Julian Gutt & Michael Klages, Alfred Wegener Institute for Polar and Marine Research, Columbus-
straoe, 0-2850 Bremerhaven, Federal Republic of Germany.

Introduction damaged during capture. The original description was

made on the basis of deep frozen material and the original
In the course of benthos investigations in the Weddell Sea colouration was visible only in the holotype. Several
(Antarctica) several holothurian specimens were photo- carmine-red spots of a diameter of approximately 10 mm
graphed and videotaped in situ. They are apparently were present on the translucent-white dorsum. Some
closely related to Bathyplotes rubipunctatus Gutt, 1990 hours after transfer of the holotype into 4% buffered
but show differences in external characteristics, not de- formaldehyde and later into alcohol, the distinctive
tectable in preserved material. This study aims to describe colouration turned into pale pink, as in the paratypes.
these phenotypes in detail, and to discuss the results This new species was only photographed once in situ
obtained by underwater photography and video from a before the original description. The photograph was
taxonomic point of view. made in the course of a longer series of benthos photo-
The results of the above mentioned faunistic survey of graphs on the shelf of the Weddell Sea. The live specimen
the distribution and ecology of holothurians in the Wed- showed a high degree of similarity with the preserved
dell Sea were recently published (Gutt in press). During holotype, mainly in colouration. Therefore it was identi-
these investigations a large number of aspidochirote fied as a representative of Bathyplotes rubipunctatus.
specimens, belonging to five species, were collected. This During successive expeditions underwater photography
holothurian group is mostly distributed in either tropical was used in the Weddell Sea for benthos analyses. As a
and subtropical latitudes or the deep sea (Deichman 1947; consequence 13 holothurians, similar to B. rubipunctatus,
Heezen & Hollister 1971a; Pawson 1 9 8 2 ~ )Our . know- were photographed. A further six specimens were
ledge of aspidochirote sea cucumbers in Antarctic waters recorded on videotape during operation of a remotely
was poor until a few years ago. However, a new species, operated vehicle (ROV) near the sea floor. These speci-
Bathyplotes rubipunctatus, has been described (Gutt mens, while similar to B. rubipunctatus, show different
1990b),along with others (Gutt 1 9 9 0 ~ )Bathyplotes
. rubi- combinations of some characteristics, indicating different
punctatus is known to be distributed mainly on the phenotypes. Heding (1942) mentioned the difficulty in-
southern shelf of the Weddell Sea, with some additional volved in separating 19 species of this ,genus from each
records from the eastern shelf (Gutt in press). It was other and from B. natuns, out of a total of 27 species, that
recognized as separate from other species of the same he investigated. The results presented here, based on
genus by the position and number of ventral and dorsal underwater photography and video, indicate that some
tube feet and papillae, the number of tentacles, and the revision of the taxonomy and systematic status of Bathy-
shape of the ossicles, mainly in the body wall. The plotes, especially its representative B. rubipunctatus, is
description, however, was not complete, because the required.
outer layer of the body wall of most types had been
thrown off by the animals during capture. This is often
Material and methods
observed among aspidochirote holothurians (Heding
1940; Pawson 1965). When the soft and fragile fragments The photographs are from the expeditions ANT 111/3, ANT VI/3 and
of the outer body wall were present, they were generally ANTVII/4 (EPOS 3) with R/V Polarstern (Hempel 1985;Fiitterer 1988;
301 Zoologica Scripta 20
302 J. Gutt and M . Klages
Fig. 1. Stations where specimens of Bathyplotes rubipunctatus or its close relatives have been observed, either by underwater photography or by
video.
Arntz et al. 1990). Most of the stations lie on the eastern shelf of the
Weddell Sea (Fig. 1) which can exceed a depth of approximately 600 m
(Carmack & Foster 1977). Some additional stations are located on the
upper continental slope. The camera was triggered at a constant distance
from the sea floor, and therefore the size of the photographed area could
be calculated (Gutt 1988). Approximately 3300 pictures, representing
3025 m2, have been made at 58 stations. The maximum operating depth
of the underwater-camera was 1200 m. The videotapes were made by a
remotely operated vehicle (ROV) in a total of 16 stations. The distance
of the transects was estimated as 15 km long and 2 m wide. Morpho-
metric measurements of the videotaped specimens were not possible, so
only a general description of them is given.
Experience from a different study shows that the colours on the slides
are those of the captured specimens in daylight.
The length measurements of the dorsal appendages of the photo-
graphed specimens are minimum values because occasionally the thin
tips of them could not be recognized exactly.
Results
Photographed and videotaped specimens are described
from the original slides and tapes respectively. No signifi-
cant differences could be detected in the general body
shape. All specimens are approximately 7 times longer
than wide, rounded anteriorly and slightly tapering at the
posterior end. Calculating a relationship between body
width and appendage length the geometric mean (GM)
regression (Ricker 1973) was calculated because the vari-
ability in both stochastic variates is mainly inherent in the
material and little of it is due to error of measurement.
The correlation coefficient for the functional regression
between the length and width of all specimens is r = 0.93
( p 5 0.05). The linear geometric mean regression model
was fitted using the ordinary least-square method to
estimate the relation between body width ( y ) and body
length (x): y = 0 . 1 5 ~+ 0.39. The significant correlation
indicates that this relationship between width and length
is similar for all measured specimens.
Chief differences, however, exist in the shape and
length of the dorsal appendages and in the colouration of
the animals. The dorsal structures can be developed as
either slight, poorly recognizable elevations of the skin or
as long filamentous appendages (see below). Apparently
intermediate types exist; nevertheless an artificial classifi-
cation of the appendages was made to combine similar
specimens into the following groups: (1) long append-
Zoologica Scripta 20
- 740
- 76’
-
50
ages, longer than the maximum body width; (2) short
appendages, shorter than body width, but at least three
times longer than their diameter at their base; (3) conical
papillae, shorter than the short appendages. It is difficult
to differentiate between the latter two categories, be-
cause it is hard to measure the diameter of the appendages
at the base. For the purpose of the present study, it is
more informative to separate these different types of
appendages or papillae in this manner than to consider
them all to be equal. The distribution of the appendages
on the dorsum is irregular, with a tendency for the longer
appendages to accumulate in the radii. The relationships
between the body width and maximum length of dorsal
appendages for different phenotypes are shown in Fig. 2.
Because of the low number of specimens no regression
model was calculated within each group. Nevertheless, a
difference between individuals of group A, together with
groups E and D and the groups B and C is intimated in the
figure..
Colouration of the entire dorsal body surface of the
specimens (see Fig. 3) is the second characteristic which
led to the classification shown in Fig. 2 and described
below. This can range from completely white with intense
red dots at the base of the longer appendages to a nearly
reddish to brownish. The diameter of the dots ranges
from 0.8 to 1.3 cm, but there is no sharp border relative to
the rest of the paler body surface. The long and short
appendages are translucent-white except at their bases
7
Icml
6
length of 5
dorsal
appendages 4
3
2
1 @E e
0
5 10 15 20 25 30
body lenglh [cm]
Fig. 2. Classification of specimens of Bathyplotes rubipunctatus or its
close relatives according to their colouration and the relation between
their body length and length of dorsal appendages, body length-body
width relation included.
 Observations on Bathyplotes (Holothuroidea) in Antarctica 303

Fig. 3 . 4 . Specimen No. 1of group A.-b. Specimen No. 7 of group A,+. Specimen No. 10 of group B . d . Specimen No. 14 of group C . 4 .
Specimen No. 15 of group C.-f. Specimen No. 18 of group D. For sizes see Table I.

(see above) and the very short ones are completely red. body length (counted in the middle of the dorsum), few
The papillae are roseate to red or brown. short appendages of transitional size between the above-
The characteristics and locations of the single speci- mentioned appendages and conical papillae. Dorsum
mens are listed in Table I. For the proposed five groups, roseate to light roseate along the translucent longitudinal
representing different phenotypes, the following general muscular structures in the radii, with approximately 8
diagnoses can be given: intense carmine-red spots at the base of most of the dorsal
appendages.
Group A . 7 specimens (Nos 1-7; Fig. 3a, b) with 5-8 short
dorsal appendages, alternating mainly in the radii, 2-5
dorsal, conical papillae per cm body length (counted in Group C. 2 specimens (Nos 14, 15; Fig. 3d, e) with 7-9
the middle of the dorsum), with a translucent white long dorsal appendages, 3-5 dorsal conical papillae per
dorsum with the exception of 5-8 intense carmine-red cm body length (counted in the middle of the dorsum),
spots at the base of most of the dorsal appendages. few appendages of transitional size between above-
mentioned appendages and conical papillae. Dorsum red,
Group B. 4 specimens (Nos 10-13; Fig. 3c) with 8-10 long brownish-red or reddish-purple with slightly more intense
dorsal appendages, 2-6 dorsal conical papillae per cm colouration at the base of most of the longer appendages.
Zoologica Scripta 20
304 J . Gutt and M . Kluges

Group D. 2 specimens (Nos 17, 18; Fig. 3f) with short
dorsal appendages, additionally approximately 6 conical
papillae per cm body length, appendages of transitional
size between above-mentioned appendages and conical
papillae, dorsum red, slightly deeper at the base of most
of the appendages (No. 18) or evenly black-reddish (No.
17).
Group E. 1 specimen (No. 19) with approximately 6 8
short dorsal appendages, significantly shorter than body
width, no additional papillae, entire body brown.
The longest appendages of specimen No. 7 (Fig. 3b) are
nearly as long as the maximum body width, therefore this
individual occupies an intermediate position. It was
classified with group A because of its colour character-
istics. The short appendages of specimens of groups A
and D are generally thicker than those of groups C and B.
No significant classification could be made for speci-
mens Nos 8 , 9 and 16 because no exact measurements of
the videotaped animals were possible. The length of
dorsal appendages of specimens recorded on videotape
(Nos 12, 13, and 17) could not be determined with the
same accuracy as photographed specimens but a classifi-
cation based on their general external morphology was
possible. The body of specimen No. 3 was not photo-
graphed completely because it was situated at the edge of
the picture.
All specimens occurred on muddy, sandy gravel with
different grain sizes. Larger pebbles (210 mm) were
present in most of the stations but distributed in patches.
Bedrock, on which sponges with a density of more than 1
specimen per m2 grow, was visible in station No. 310. The
rich sessile epifauna, well known in some areas of the
Weddell Sea shelf (see Hempel 1986; Gutt & Klindt 1988;
Gutt et al. 1988; Gutt & Vogel 1988; Gutt 1990c) is
generally missing at stations where Bathyplotes spp.
occur. Species of this genus are found on the upper
continental slope, where the epifauna covers an average
of 10% of the sea floor. Exceptions are found in the
shallower stations No. 438 and No. 396 where a maximum
of 50 or 90% of the sediment is covered by bryozoans and
hydrozoans. Here also, crinoids are abundant.
Discussion
In his faunistic study Gutt (in press) determined many
holothurians to be representatives of the recently de-
scribed Bathyplotes rubipunctatus. Only one in situ photo-
graph and damaged specimen was available. Several
characteristics were investigated in this preserved
material and no significant differences were found be-
tween the specimens. The new observations using under-
water photography and video show the presence of more
than one phenotype, which cannot be identified using
only preserved material. The length of the dorsal append-
ages is the best characteristic for separating different
groups, although transitional types exist within a broad
range.
The basic colour of the specimens ranges from com-
pletely white to pale roseate, red to red-violet, red-brown
to blackish-red. Classifications were made to keep colou-
ration constant within one group. All specimens (except
Nos 13,16 and 18) have in common a more or less intense
colouration at the base of some dorsal appendages. These
can appear as distinct carmine-red spots in the white to
roseate individuals or as slightly darker areas in the
reddish animals. Several kinds of transitional colou-
rations seem to be possible because no distinct limits
between the different types can be drawn. In other related
species, such dots are also described. Bathyplotes mose-
leyi has whitish warts (Th6el 1886), B. crenulatus and B.
variabilis have greenish ones (Koehler & Vaney 1905).
Holothuria mexicana, H. dakarensis and B. natans have
brown spots (Pawson 1968,1976;Pawson & Shirley 1977).

Table I. Characteristics and occurrence of differentphenotypes of Bathyplotes in the Weddell Sea, recorded using underwater
photography or video

No. of appendages No. of short

Cruise and Frame No. No. of Diameter longer than papillae per
No. station No. (time) dots of dots body width 5 cm body length Group

1 11113310 43 7 0.9 - approx. 12 A

2 VI13396 72 8 0.9 - approx. 10 A
3 VI13 396 60 not photographed in its complete length A
4 VII14260 1155 7 0.9-1.1 - very dense A
5 VIIl4 259 1188 * 6 0.9 - dense A
6 VII14294 325 approx. 5 approx. 0.9 ? ? A
7 VII14293 495 8 1.1-1.3 - 25 A
8 VI13 314 16:20 approx. 8 ? ? ? A or B
9 VIl3 314 16:33 approx. 8 ? ? ? A or B
10 VII14 294 328a 8 1.1 10 25 B
11 VII14 294 299b 8 ? 8 12 B
12 VI13 332 15:58 8 ? present ? B
13 VI13 332 16:32 9 ? present ? B
14 VII14294 330 8 1.3 10 22 C
15 VIIl4294 341 7 0.9 11 12 C
16 VIl3 472 13:30 - - approx. 18 ? C or D
17 VIl3 438 14:17 - - probably none dense D
18 VII14 294 356 - - - dense D
19 VIIl4294 301 - - - 6-8* E

*On the entire dorsum.

Zoologica Scripta 20
 Observations on Bathyplotes (Holothuroidea) in Antarctica
On the basis of the five phenotype-groups described
here, one might assume the presence of a theoretical
maximum of five different species. Additional infor-
mation such as microscopical investigations of the same
specimens, are needed. The main two characteristics,
number and shape of the papillae and colour, occur in a
broad range of intensity with a more or less continuous
gradient. They also appeared in different combinations in
the examined material. At present we cannot decide
whether these groups represent new species or just differ-
ing phenotypes of one species. It is possible that there are
more than three species of the genus Bathyplotes in high
Antarctic regions. The other two species are B. fuscivin-
culum Gutt, 1990 and B. moseleyi (ThCel, 1886). B.
fuscivinculum is significantly different from the specimens
described here (see Gutt 1990b). Several descriptions of
B. moseleyi (Ekman 1925,1927; Ludwig 1898) are similar
to that of B. rubipunctatus. A red to violet colour occurs
in the Aspidochirotida e.g. Stichopus roseus (Augustin
1908), B. natans (Ostergren 1896) or B. goldenhindi
(Mitsukuri 1912) as well as in the Elasipodida e.g. Ben-
thodytes sanguinolenta (Pawson 1982b). However, in all
the material caught, no specimens have been found which
were similar to those in the photographs and which could
be determined as elasipod holothurians. Only for speci-
men No. 19 does the photographed individual fit with the
general description and drawings of the Psychropotidae
(Elasipoda) made by Hansen (1975). For all other speci-
mens however, a close relation to B. rubipunctatus must
be assumed.
Concerning the general distribution of B. rubipunctatus
and its close relatives in the southern and eastern Weddell
Sea, this study agrees with Gutt (in press) although the
sample number is not high enough for statistical calcu-
lations. The main depth of distribution of the above
described individuals is the deeper part of the shelf (300-
600 m) or the upper slope (2600 m). Only seven speci-
mens were photographed on the shelf whereas only one-
third of the photographed area was situated on the upper
slope where nine specimens were found. The area investi-
gated by the ROV was approximately four times larger,
and only two of the described specimens were found. The
fact that more than one-third of all these animals are from
one station (No. 294) supports the suggestion that B.
rubipunctatus has a tendency to be distributed in patches
as already mentioned by Gutt (in press). The relationship
between the occurrence of B. rubipunctatus and soft
bottom, suggested by Gutt (in press) cannot be confirmed
by this study.
The absolute abundance of B. rubipunctatus and its
relatives can roughly be estimated as very low, with the
exception of station No. 294 which has an average of 0.2
specimens per m2. Ohta (1983)found average densities of
2 specimens per 1000 m2 for other species of the genus
Bathyplotes and a maximum value of 30 specimens per
1000 m2 for Bathyplotes goldenhindi off Japan in depths
between 295 and 539 m by means of underwater photo-
graphy.
The present results show that neither underwater pho-
tography nor the investigation of preserved material
alone leads to a satisfactory conclusion in specific taxono-
mic investigations. For future ecological and taxonomic
305
studies, more direct observations and published photo-
graphs of deep sea holothurians like those of Menzies
(1963),Pawson (1982b) and Heezen & Hollister (1971b)
are necessary for new descriptions, as well as for the
confirming identification of specimens belonging to
already known species.
Acknowledgements
We are grateful to Alodia Holierhoek for critical review of the manu-
script. Data presented here were mainly collected during the European
“Polarstern” Study (EPOS) sponsored by the European Science Foun-
dation and the Alfred Wegener Institute for Polar and Marine Research.
This is publication No. 263 of the Alfred Wegener Institute.
References
Arntz, W., Ernst, W. & Hempel, I. 1990. The Expedition ANTARK-
TIS VII/4 (EPOS 3) and VII/S of “Polarstern” in 1989.-Ber.
Polarforsch: 1-214.
Augustin, E. 1908. Uber japanische Seewa1zen.-Abh. buyer. Acad.
Wiss. Math-phys. K1. 2: 1 4 4 .
Carmack, E. C. & Foster, T. D. 1977. Water masses and circulation in
the Weddell Sea. In Polar oceans proceedings of the polar oceans
conference, Montreal, May 1974 (ed. M. J. Dunbar): 167-177. Arctic
Institute of North America, Calgary.
Deichman, E. 1947. Shallow water holothurians from Cab0 de Hornos
and adjected waters.-An. .Mus. argent. Cient. nut. 42: 325-351.
Ekman, S. 1925. Ho1othurien.-Further zool. Results Swed. Antart.
Exped. 1901-1903 I: 1-194.
Ekman, S. 1927. Holothurien aus der Ostantarktis und von den
Kergue1en.-Dt. Siidpo1.-Exped. 1901-03 (Zool.) 11: 359419.
Fiitterer, D. K. 1988. Expedition ANTARKTIS-VI mit FS “Polarstern”
in 1987/1988.-Ber. Polarforsch. 58: 1-267.
Gutt, J. 1988. Zur Verbreitung und okologie der Seegurken (Holothur-
oidea, Echinodermata) in Weddellmeer (Antark&).-Ber. Polar-
forsch. 41: 1-87.
Gutt, J. 1990a. New Antarctic holothurians (Echinodermata)-I. Five
new species with four new genera of the order Dendrochir0tida.-
2001.Scr. 19: 101-117.
Gutt, J. 1990b. New Antarctic holothurians (Echin0dermata)-11. Four
species of the orders Aspidochirotida, Elasipodida and Apodida.-
Zool. Scr. 19: 119-127.
Gutt, J. 1990c. 3.9 Underwater photography. In The Expedition
ANTARKTIS VIV4 (EPOS 3) and VII/5 of “Polarstern” in 1989.-
Ber. Polarforsch. 68: 107-110.
Gutt, J. in press. On the distribution and ecology of holothurians in the
Weddell Sea (Antarctica).-Polar biol.
Gutt, J., Gerdes, D. & Klages, M. 1988. Benthic fauna.-Biomass
newsl. 10: 10-11.
Gutt, J. & Klindt, H. 1988. Direktbeobachtungen an benthischen
Besied1ungsstrukturen.-Ber. Polarforsh. 58: 100-105.
Gutt, J. & Vogel, S. 1988. Leben im eisigen Dunkel. “Polarstern”-
Expedition 1988.-Spektrurn Videothek.
Hansen, B. 1975. Systematics and biology of the deep-sea holothurians.
Part 1. E1asipoda.-Galathea Rep. 13: 1-262.
Heding, S. 1940. Die Holothurien der deutschen Tiefsee-Expedition. 11.
Aspidochirote und Elasipode Formen.-Wiss. Erebn. - dt. Tiefsee-
E&d. 24: 319-375.
Hedine. S. 1942. Holothuroidea Part 11. AsDidochirota-ElasiDoda-
D&drochirota.-The Danish Zngolf-Expedition. Vol. IV: 1-34.
Heezen, B. C. & Hollister, C. D. 1971a. The deep, deep sea floor.-
Nut. Hist. 80: 30-33.
Heezen, B. C. & Hollister, C. D. 1971b. Face of the deep. Oxford
University Press, Oxford.
Hempel, G. 1985. Die Expedition Antarktis 111mit FS “Polarstern”.-
Ber. Polarforsch. 25: 1-223.
Hempel, G. 1986. Funf Jahre Schwerpunktprogramm “Antarktisfors-
chung” der Deutschen Forschungsgemeinschaft Ruckblick und
Ausb1ick.-Ber. Polarforsch. 29: 1-150.
Koehler, R. & Vaney, C. 1905. Echinoderma of the Indian Museum:
account of the Deep-sea Holothurioidea collected by the R.I.M.S.
Investigator (ed. A. Alock), Calcutta: 1-123. Friedlander & Sohn,
Berlin.
Ludwig, H. 1898. Ho1othurien.-Ergebnisse der Hamburger Magalhae-
nischen Sammelreise 1892193, I. Band: 1-98.
Zoologica Scripta 20
306 J . Gutt and M . Klages
Menzies, R. J. 1963. General results of biological investigations on the
deep-sea fauna made on the U.S.N.S. Eltanin (U.S.A.R.P.) during
cruise 3 between Panama and Valparaiso, Chile in 1 9 6 2 .4 n t . Rev.
Ges. Hydrobiol. 48: 185-200.
Mitsukuri, K. 1912. Studies on actinopodus Ho1othuroidea.-J. Coll.
Sci. imp. Univ. Tokyo 29: 1-284.
Ohta, S. 1963. Photographic census of large-sized benthic organisms in
the bathyal zone of Suruga Bay, central Japan.-Bull. Ocean Res.
Inst. Univ. Tokyo 15: 1-244.
Ostergren, H. 1896. Zur Kenntnis der Subfamilie Synallactinae untes
den Aspidochiroten. In Zoologiska studien-festskrift Wilhelm Lillje-
borg tillegnad pb hans dfjionde fodelsedag af svenska zoologer: 345-
360. Almquist & Wiksell, Uppsala.
Pawson, D. L. 1965. The bathyal holothurians of the New Zealand
region.-Zoology Publs Vict. Univ. Wellington 39: 1-33.
Pawson, D. L. 1968. The echinozoan fauna of the New Zealand
Subantarctic Islands, Macquarie Islands, and the Chatham Rise.-
N . Z . Dept. Scient. Industr. Res. Bull. 187: 1-35.
Zoologira Srripta 20
Pawson, D. L. 1976. Shallow-water sea cucumbers (Echinodermata:
Holothuroidea) from Carrie Bow Cay, Belize.-Proc. biol. SOC.
Wash. 89: 369-382.
Pawson, D. L. 1982a. Holothuroidea. In Synopsis and classifcation of
living organisms (ed. S. P. Parker): 791-818. McGraw-Hill, New
York.
Pawson, D. L. 19826. Deep-sea echinoderms in the tongue of the ocean,
Bahama Islands: a survey, using the research submersible Alvin.-
Mem. Aust. Mus. 16: 129-145.
Pawson, D. L. & Shirley, T. C. 1977. Occurrence of the subgenus
Holothuria (Holothuria) in the Gulf of Mexico (Echinodermata:
Holothuroidea).-Proc. biol. SOC.Wash. 90: 915-920.
Ricker, W. E. 1973. Linear regression in fishery research.-.I. Fish. Res.
Bd Can. 30: 409434.
ThCel, H. 1886. Report on the Holothuroidea Part 2.-"Challenger"
Sci. Results-Zoology 14: 1-290.