A new species of Biemna (Porifera, Poecilosclerida) from the Sultanate of Oman

10 Pages·2002·2.3 MB·English

by  Van Soest, R W M

Beaufortia
BULLETIN ZOOLOGICAL MUSEUM
UNIVERSITYOF AMSTERDAM
Vol. 52, no. 2 July 26, 2002
A new species of Biemna (Porifera, Poecilosclerida) from the
Sultanateof Oman
R.W.M. vanSoest & E.J. Beglinger
InstituteforBiodiversity andEcosystem Dynamics (£odlogisch Museum), University ofAmsterdam,
P.O. Box 94766, 1090 GTAmsterdam,Netherlands
[email protected], [email protected]
Abstract
Recent(1998)collectingoffthe coastsofOmanin thecourseof theEC-MAS3 funded‘Symbiosponge’ projectyieldedthree
species of the genus Biemna Gray, 1867 (Demospongiae, Poecilosclerida, Mycalina, Desmacellidae) notpreviously known
from the area.One ofthese appeared tobe newtoscience and is describedasBiemna omanensisn.sp..Itsdistinguishingfea-
tures comprise the possession of oxeamegascleres, which is of rare occurrencein Biemna, and a full complement of
microscleresincluding microxeas, sigmas, commata, all of which aredivisiblein two non-overlapping size categories, and
raphides. One ofthe other twospecies conformedtoanearlier described species Biemna ciocalyptoides Burton, 1959,which
was discoveredtobe ajuniorsynonym ofBiemna ehrenbergi(Keller, 1889)(new combination).The third species is the com-
monand widespreadBiemna tubutataDendy, 1905.
Keywords:Porifera, Oman,Biemna,new species
INTRODUCTION Murray Expedition, 1933-34, and reported by
Burton(1959).
The Sultanateof Oman occupies the southeast In the course of the EC-MAS3 'Symbio-
corner of the Arabian Peninsula and is located sponge' project, sponges were collected inseveral
between latitudes 16°40'and 26°20'Nand longi- localities along the coast of Oman for the pur-
tudes 51°50'and 59°40'E. Thecoastline extends pose of detecting novel bioactive compounds.
1.700 km, from the Strait of Hormuz in the Several of these sponges appear new to science
north, to the Republic of Yemen in the south, and in an ongoing series of publications we will
andborders onthreemarineregions: theArabian describe these. The present study deals with a
Gulf, Gulf of Oman and theArabianSea. From newspecies ofBiemna, and also includes descrip-
the Oman coasts about 100 species of sponges tions of two other Biemna species already known
have beenrecorded, and an additional30 species from nearby areas, but not yet from Oman
are known from nearby South Arabian localities. waters.
Mostof thesesponges were collectedby theJohn The genus Biemna is recognizable among
15
Fig. 1.Mapofthe Sultanate of Oman.Localities ofBiemnaspecies areindicatedby*. 1 =B.omanensis n.sp.,2 =B.ehrenber-
gi,3 =B.tubulata.
Desmacellidae on the combination of microscle- of tylostyles. Biemna is represented in the Indian
res (sigmas, microxeas, commata,raphides) and a Ocean by approximately 12 species (review in
confusedto plumose arrangement of the megas- Hooper, 1996). Some of these records from the
cleres (Hajdu & VanSoest, 2002). Closely related western Indian Ocean concern species with
generaareNeofibularia withareticulate skeletonof allegedly wide Indo-Pacific distributions, but may
strongyles, andDesmacellawithaplumose skeleton eventually turn out to benew taxa.
16
MATERIALSANDMETHODS DESCRIPTION. - Thick cushion, of approx. 10 cm
diameter and thickness (Fig. 2A) (the actual
Specimens studied were collected using SCUBA retainedholotype is a large fragmentof this spec-
and snorkeling by Raquel Gomez andRobert G. imen, the largest part having been sacrificed for
Moolenbeek, Zoological Museum of Amsterdam chemical investigations ). Surface with numerous
(ZMA) in November 1998. Collecting localities irregular projections and villi of approx. 1.5 cm
are given inFig. 1. Specimens were photograph- high. Surface inbetween thevilli covered insedi-
ed insitu or on deck beforepreservation in 70% ment and algae, leaving small oscules visible.
alcohol. Subsamples were collected for testing of Colour yellow, but sediment covered parts grey-
biological activity andfor cell biological observa- ish. Consistency soft, easily crumbled. Touching
tions.Preserved sampleswere incorporated in the with bare hands causes severe burning sensation
collections of the Zoological Museum of and prolonged itching.
Amsterdam. For comparison, Biemna material Skeleton: basically alveolar to confused, with
from the ZMA collections (Red Sea, Seychelles, many spicules and microscleres strewn without
Curasao) was consulted, and a slide made from direction. Parallel tracts of 5-10 spicules thick
type material of the HumboldtMuseum, Berlin support the villi.Surface skeleton mostly absent,
(ZMB) was examined, obtainedwith permission but in the surface membranethereis a discontin-
from thecurator Dr D. Kuhlmann, now retired. uous cover of microxeas and sigmas carried by
For microscopical studies, tangential and per- projecting tracts of megascleres. Microxeas and
pendicular sections were made with a scalpel; other microscleres are also very common in the
these were dried on a hotplate, subsequently interior.
mountedinCanadabalsam, andexaminedusing Spicules: megascleres oxeas (Fig. 2B), smooth,
100-400x magnification. Furthermore, a frag- symmetrically curved, sharply pointed, 230-
ment of approx. 3 mm3 was cooked in concen- 255.7-286 x 7-8.3-10 (am. Microscleres microx-
trated HNO3, the residue was washedand cen- eas, sigmas, and commata, and raphides, all of
trifugated 5 timesin distilled water,and suspend- which may occur in dragmata. Microxeas,
ed in 96% alcohol; spicule suspensions were straight, rugose/microspined, in two non-over-
pipetted on SEM stubs and on microscopic glass lapping size categories: 123-140.4-147x 2-2.9-4
slides; the dried spicules were mounted in jxm (Fig. 2CJ-C2) and 63-75.5-87 x 2-2.8-4
Canada balsam for light microscopy and sput- (Fig. 2D|-D2). Thin sigmas occur in two non-
teredwith gold for examinationunderSEM. overlapping size categories, 27-31.6-37|j.m (Fig.
Spicule size data are based on 25 measure- 2E) and 10-12.1-14 (Fig. 2F). Commata in
ments for eachspicule category. two non-overlapping size categories, 27-37.5-51
|im (Fig. 2Gi"G2) and 14-16.8-18 (im (Fig. 2H).
Raphides (Fig. 21), thin, possibly thin growth
SYSTEMATIC DESCRIPTIONS
stages of the larger microxeas, 112-123.2-135x
0.5 fim.
Class Demospongiae
OrderPoecilosclerida
DISTRIBUTION.- Knownonly fromthetype local-
SuborderMycalina
ity, N point Masirah Island, Oman, on hard sub-
Family Desmacellidae
strate in sandy environment.
Genus BiemnaGray, 1867
BlOACTIVlTY. - The sponge methanolic extract
Biemnaomanensis n.sp.
was found to be mildly active against brine
Figs. 2A-I
shrimp larvae andseveral parasitic fungi.
MATERIAL. - Holotype: ZMAPOR. 14640,Sultanate of ETYMOLOGY. - Named after the Sultanate of
Oman, Masirah Island, N point, 20.70°N 58.78°E (dec.
Oman.
coord.), onwreck of S.S. 'Electra' and on nearby stones,
depth 7.6 m,98/IO/NOV15/RG/046,coll. R. Gomez &
R.G. Moolenbeek, 15-XI-1998. DISCUSSION. - No species of Biemna have previ-
17
Fig.2.Biemna omanensisn.sp. A,photoofhabittaken immediatelyaftercollection.B-I, SEMphotosofspicules,size barsindi-
cate 100 μm (B), 10μm (C1,D1, E1, F, G1,H), and 1 μm (C2, D2, E2, G2). B, oxea.C, large microxea. D, small microxea.
E,large sigma.F, small sigma. G, largecomma.H,small comma. I, raphide.
ouslybeenreported fromOmanwaters.From the recorded two species of Biemna, B. ciocalyptoides
Gulf of Aden at the opposite end of the south Burton, 1959 from 11°57'12"N50°35'E, 37 m,
coast of the Arabian peninsula, Burton (1959) and B. fortis (Topsent, 1897) from two localities:
18
11°53'42"N 51°13'12"E, 73-220 m, and 11°57' havecrowns of spines on theircommata.
12"N 50°35'E, 37 m. The former species is a A furtherfeature revealedby SEMand worthy
junior synonym of B. ehrenbergi (Keller, 1889, see of emphasis are the roughened ends of the sig-
below). Both species have styles for megascleres, mas inthis genus(Fig. 2E2).Thisfeatureis shared
bothhavemuch larger megascleres (up to 1000x withNeofibularia (e.g. Van Soest, 1984).
30 (am) and lack commata. Below we record the
occurrence ofB. tubulatainOmanwaters; thishas Biemnaehrenberei (Keller, 1889) n. comb.
a tubular habitand smaller microxeas. Figs. 3A-G
Hooper (1996) provides an overview of all
Biemna species recorded from the Indian Ocean. Acanthella ehrenbergi Keller, 1889: 395.
Noneof thempossess oxeas as megascleres. Just Biemnaciocalyptoides Burton, 1959:227, text-fig. 14
outside the Indian Ocean, from Philippine (not: Sigmaxinella ciocalyptoides Dencly, 1897:242;
waters, an oxea-bearing species was reported, nec Biemna (Allantophora) ciocalyptoides; Hall-
Biemna gellioides Levi & Levi, 1989. This species mann, 1916:514, pi. XXX figs 6-8).
has much larger oxeas (500-600 jam); microxeas
are drawn(Levi&Levi, 1989:fig. 46.3), butinthe MATERIAL. - ZMA POR. 14641, Sultanate of Oman,
text theseare calledraphides, and they are clear- Masirah Island, NE point, (dec. coord.)20.70°N 58.90°E,
ly much smaller than those of B. omanensisn.sp.; depth 11.8m, 98/IO/NOV15/RG/047,coll.R. Gomez&
R.G.Moolenbeek, 15-XI-1998.
sigmas are similar to those of B. omanensis n.sp.,
butcommata are apparently lacking.
Elsewhere, in the Caribbean, a Biemna with DESCRIFI'ION. - From a massive base buried in
oxeas isalso found, B.cribaria(Alcolado & Gotera, the sandy bottom arise cylindrical to conical fis-
1986), with synonym B. oxeata Van Soest & tules (Fig. 3A) of5-6 cm length, 1-2 cm in thick-
Stentoft, 1988. This has larger oxeas (300-550 ness,with thesurface provided withcharacteristic
fim), larger sigmas (50-77 and 20-26 jam), and longitudinal, somewhatcorrugated, grooves. The
raphides in two size categories (50-80 and 10-20 fistules end in one or more thin projections.
jam). No commata or rugose microxeas are Colour noted as mustard-brown in life, pale
reported. brown inalcohol. Consistency firm to hard.
Thedermatitisproduced by thisspongeis rem- Skeleton: axially compressed in the fistules, with
iniscent of Biemna laboutei Hooper, 1996 and is thick columnsof megascleres, 10-20in cross sec-
also foundin several otherspecies of Biemna and tion, ascending longitudinally and extra-axial
the closely related Neofibularia. B. laboutei from thinnerbundles and single megascleres directed
Madagascar is a ramose orange species, with towards the surface. In the main body, spicules
styles as megascleres, lacking commata, and thus are arranged in confusion. Spongin is visible in
clearly differs from our newspecies. variable quantities. Numerous microscleres are
Figs 2G]-G2 and 2H (as well as Figs 4G-FI of strewn throughout the interiorand in thesurface
B. tubulata) are the first SEM images made of membrane.
commata of Desmacellidae. Previously, only Spicules: styles (Fig. 3B|-B2), smooth, robust,
thoseof Merlia(Merliidae) were examined under curved, 596-740.0-806 x 21-27.9-33 jim.
SEMandlike thoseof theBiemnacommata, these Microxeas (Fig. 3C), often in dragmas, thin,
are microspined/rugose. A new and apparently raphide-like: 36-41.0-45 x 1 |jm; sigmas, often in
unique feature for Biemna commata is the crown sigmodragmas, in three size categories, 54-61.6-
of spines found on the heads. Under light 66 (im (Fig. 3D), 21-29.0-35 (Fig. 3E), and 13-
microscopy, such adetailis onlybarely visibleand 13.8-15 |-im (Fig. 3F); trichodragmas, flexuous,
so farhas escaped observationby spongetaxono- 240-270 x 13-30 fim, individual raphides (Fig.
mists. This ornamentation is here regarded as a 3G) 177-197.3-215x0.5 p.
synapomorphy forBiemnaspecies possessing com-
mata, since both the present species and B. tubu- BIOACTIVITY.- Themethanolicextractwas found
lataappeartopossess similaradornment. Itispre- to be mildly active against brine shrimp larvae
dicted thatcommata of otherBiemnaspecies also andseveral species of parasitic fungi.
19
Fig. 3. Biemna ehrenbergi(Keller, 1889).A, photo ofhabit taken immediately aftercollection. B-G, SEMphotos of spicules,
size bars indicate 100 μm(B1, G), 10μm (B2, C, D1, E,F), and 1 μm(D2).B, style. C, microxea. D, largesigma. E, middle
sigma.F, small sigma. G, raphide.
DISTRIBUTION. - Red Sea, South Arabia, nearby Gulf of Aden. This is stated to have a
Seychelles. massive pyramidal base with numerous fistular,
aculeateprocesses, which adequately conformsto
DISCUSSION. - We were able to confirm the spe- the habitof the Omanmaterial. The spicules of
cific identity of our materialwithAcanthella ehren- Burton's specimen appear very similar in shape
bergiKeller, 1889by examinationof aslide of the and size, the differences being a slightly smaller
ZMB holotype. Ourmaterial is closely similar to size of the styles, the presence of microxeas of
Biemnaciocalyptoides Burton, 1959, described from only 40 jam, and trichodragmas not exceeding
20
270 jam. The microxeas mentioned by Burton In dry condition, the two specimens are cited as
measured 160 fim, and this may conceivably be black. Thiscontrasts withthe beige to yellow-grey
straight raphides from the trichodragmas which colour in alcohol reported for the Southern
were observed to be that size, as the distinction Arabian material. However, in the Seychelles,
between microxeas and raphides may be difficult where the species appears common (see Van
to assess in dissociated spicule mounts. The Soest, 1993as Biemna ciocalyptoides sensu Burton),
absenceof thesmallmicroxeasandthelonger tri- colours of live specimens and alcohol specimens
chodragmas remain discrepancies of some con- varied from pale brown to dark wine-red.
cern, but it is proposed here to consider Burton's Apparently, colours are variableforreasons so far
sample conspecific with ours. unknown.All thesecombinedsimilarities suggest
As Hooper (1996) pointed out, Biemna ciocalyp- thatthe SouthArabianmaterialdescribedabove
toides was already in use for the combination is best considered Biemna ehrenbergi until more
Biemna ciocalyptoides sensu Dendy, 1897 (as detailed analysis of population differences
Sigmaxinella). Dendy provided no illustrations, but between Red Sea, Arabian Sea and the
the habit was extensively redescribed by Seychelles is made.
Hallmann, 1916, who included several photos. Hooper (1996) suggested thatBiemna seychellen-
From thesedescriptions, itis clear that it is simi- sis Thomas, 1973is conspecific with this species
lar in growth form to the South Arabianspecies, (that is: with B. ciocalyptoides sensu Burton), but
but differs substantially in spicule dimensions: Thomas described commata in his material,
styles are clearly smaller (500 x 18 jam), sigmas whichare absentfrom B. ehernbergi. The length of
occur only in a very small category (17 (am), and the trichodragmas is given as 28-48 |am only,
trichodragmas were cited as 30 (am. The latter which contrasts with the characteristic tri-
may have includedmicroxeas, of whichno men- chodragmas of 160+ |j.min B. ehrenbergi. As faras
tion is made, however. Clearly, there is no match thehabit can be deducedfromThomas' descrip-
in these spicule sizes and categories with South tion of the dried material, this appears also dif-
Arabian specimens, andaccordingly - andin con- ferent.
junction with Hooper, 1996 - we conclude that
thetwo are to beregarded as distinct species. Biemnatubulata(Dendy, 1905)
Fortunately, aseniorsynonymof B.ciocalyptoides Figs. 4A-I
sensu Burton is available in Biemna ehrenbergi
(Keller, 1889 as Acanthella). The existence of an Desmacella tubulataDendy, 1905: 155, pi. IX fig. 4.
earlier name for this species is here reported for ?Biemna microxa Hentschel, 1911: 316, fig. 15.
the first time. Keller (1889) described Acanthella Toxemna tubulata; Hallmann, 1917:673.
ehrenbergi in doubtful terms and also omitted to ?Biemnaanisotoxa; Vacelet& Vasseur, 1971:90, fig.
add an illustration, which explains why a clear 42 (not: B. anisotoxa Levi, 1963).
assignment to its properplace in the spongeclas-
sification was lacking so far. Reexaminationof a MATERIAL - ZMA POR. 14604, Sultanate of Oman,
slide made from the ZMB holotype, nr. 157, Hramal Road, near government building, (dec. coord.)
revealed that large numbers of sigmas and 23.57°N 58.62°E, 8 m, between stones in sandy environ-
ment, 98/IO/NOV06 /RG/008, coll. R. Gomez & R.G.
raphides supplement theskeleton ofrobuststyles.
Moolenbeek 6-X1-1998.
The spiculation comprises styles, commonly also
stair-stepped oxeote modifications of 500-700 x DESCRIPTION. - Groups of thin-walledtubes (Fig.
20 sigmas I 55-60 fj.m, sigmas II 20-28 |am , 4A), 3-5 cm high, 1.5 cm diameter, with vents
sigmas III: 12 |am; trichodragmas 220-250 x 30- approx. 1 cm in diameter, growing from a base
65 (im, individual raphides shorter, of variable lodged betweenandundersmall stones and octo-
length, hair-thin(microxeas are not discernibleas corals. Partof thebodymay consistoflobateout-
aseparatespicule category). ThusA. ehrenbergiis a growths. Surface microconulose, slighdy rough,
clear Biemna. Its habitis describedas 'massive pil- oftenencrustedwith sandgrains. Colour: orange-
lar-shaped' with on the upperside finger-shaped yellow in life, cream to light beige in alcohol.
projections of several cm high and 1.5 cm thick. Consistency very soft, easily torn.
21
Fig. 4.Biemna tubulata (Dendy, 1905).A,photoofhabit taken in situ.B-I, SEMphotosofspicules,size bars indicate 100 μm
(B), 10μm(C, D, E1,F, G, H and I), and 1 μm(E2).B, style. C, largemicroxea. D, small microxea. E, largesigma. F,small
sigma. G, large comma.H,small comma. I,raphide.
Skeleton: surface membrane carried by an ill- be formed by protruding megascleres and thick
defined tangential skeleton made up of single bundles of raphides. The choanosomal skeleton
megascleres and a dense mass of raphides. In is made up of a lax, confused arrangement of
cross section, the surface conules are observedto megascleres, without visible consolidating spon-
22
gin, andaprofusion of raphides and sigmas. were not mentionedby Hentschel. The shape of
Spicules: thin, smooth, curved styles (Fig. 4B), the sponge is given as 'massive withwide canals'.
295-306.2-324 x 4-5.2-6 pm, microxeas in two The species was also recorded from the West
non-overlapping size categories, 75-88.8-114 x Indian region (e.g., Van Soest, 1984: 133), but
1.5-2.1-3.5pm (Fig. 4C), and 21-28.4-37 x 1 nm that hassubsequently beenassigned to aseparate
(Fig. 4D); thin sigmas two size categories, which species, B. caribeaPulitzer-Finali, 1986, because it
appearto be almost overlapping, 24-30.6-39pm differs in having a less clearly tubular habit.
(Fig. 4E), and 14-17.1-21 pm (Fig. 4F); commata Spicule sizes and categories are remarkably simi-
in two non-overlapping size categories, 28-36.2- lar, but the smaller microxea category differs by
45pm(Fig. 4G), and 12-14.1-18jam(Fig. 4H), tri- being clearly shorter inB. tubulata.
chodragmas, 135-160.4-180 x 18-31.0-45 pm,
individualraphides (Fig. 41), 91-105.3-123x 0.5 ACKNOWLEDGEMENTS
pm.
Oman sponges were collectedandphotographed
BIOACTIVITY. - The rough methanolic extract by Raquel Gomez and Robert G. Moolenbeek.
was not active against test organisms, but one of This study as well as the fieldwork was financed
the further refined fractions was strongly active under grant EC-MAS3-CT97-0144. Dr Adil
against brineshrimp larvae. Mohammed A1 Qasim of the Marine Science
and Fisheries CentreinMuscatis thankedfor the
DISTRIBUTION. - Sri Lanka, India, Seychelles, use of facilities duringthe stayatMuscat. Martin
Providence (Indian Ocean), South Arabia, SW Day (RAF-O) provided SCUBA facilities on
Australia. Masirah.
DISCUSSION. - This species has been recorded REFERENCES
from allcorners of theIndianOcean. Compared
to the original description of Dendy from Sri ALCOLADO,P. M.& G.G. GOTERA, 1986. Nuevas adi-
Lanka, our specimen differs inhaving commata, cionesala fauna deporiferos de Cuba. Poeyana 331: 1-
notmentionedby Dendy. However, Dendy (1916) 19.
reported 'toxas'of 29-32 (im, whichcould repre- BURTON, M. 1959. Sponges. In: Scientific Reports. John
MurrayExpedition, 1933-34. 10 (5): 151-281.
sent the commata.
DENDY, A. 1897. Catalogue of Non-Calcareous sponges
Biemna fistulosa (Topsent, 1897 as Desmacella) collected by J. Bracebridge Wilson, Esq., M.A., in the
from Indonesiais reminiscent of this species, but neighbourhoodof PortPhillip Heads. Part III.Proc. R.
differs inhaving sigmas of upto 60 jam, clearly in Soc. Victoria (n.s.) 9: 230-259.
excess ofthe sigmasof thetype andof our mate- DENDY, A. 1905. Report on the sponges collected by
rial. Professor Herdman,atCeylon, in 1902. In: Herdman,
Biemna anisotoxa sensu Vacelet & Vasseur, 1971 W.A. (ed.). Report totheGovernment of Ceylon onthe
Pearl Oyster Fisheriesof the Gulf ofManaar. 3 (suppl.
fromMadagascar couldvery well bespecimens of 18):57-246.
this species, as theirshape is tubular, andtheonly DENDY, A. 1916. Report onthe non-Calcareous sponges
difference in the spicule complement of their collected by Mr. James Hornell at Okhamandal in
specimens is the lack of differentiatedsize cate- Kattiawarin 1905-06. Rept. Government ofBaroda on
the Marine Zool. ofOkhamandal in Kattiawar. Part II:
gories of commata (called 'microstyles toxi-
93-146.
formes'). Levi's (1963) type from South Africa GRAY, J. E. 1867. Notes on the arrangementofsponges,
appears different. It has three size categories of with the descriptions of some new genera. Proc. zool.
sigmas and lacks tubes, so this remains a valid Soc. London 1867: 492-558.
species. HAJDU, E.& R. W. M. VANSOEST, 2002 (in thepress).
Hooper (1996) suggested that Hentschel's Family Desmacellidae. In: Hooper,J. N. A.& R. W.M.
(1911) B. microxa from SW Australia is a junior van Soest (eds). Systema Porifera. Kluwer Academic/
PlenumPress, New York, London.
synonym of B. tubulata. No conclusion can be HALLMANN, E. E 1916. A revision of the genera with
drawn until the type has been reexamined and microscleres included, or provisionally included, in the
the presence of commata is established, as these family Axinellidae; with descriptions of someAustralian
23
species. Part II. Proc.Linn. Soc. N. S.W. 41(163): 495- PULITZER-FINALI,G. 1986.A collection ofWestIndian
552. Demospongiae (Porifera). In appendix, a list of the
HALLMANN,E. F. 1917 [1916]. Arevision ofthe genera Demospongiae hitherto recorded from the West Indies.
with microscleres included, or provisionally included,in Ann. Mus.Civ. Storia Nat. Genova 86: 65-216.
the family Axinellidae; with descriptions of some SOEST,R.W.M.VAN 1984.Marine spongesfromCurasao
Australianspecies. Part III.Proc.Linn. Soc. N. S.W. 41 and other Caribbean localities. Part III.Poecilosclerida.
(164): 634-675. Stud.Fauna CurasaoCarib. Isl. 66 (199): 1-167.
HENTSCHEL,E. 1911.Tetraxonida.2.In: Michaelsen,W. SOEST,R.W.M.VAN 1993. Sponges ofthe Seychelles. In:
& R. Hartmeyer(eds). Die Fauna Siidwest-Australiens 3 J. van der Land (ed.). Oceanic reefs of the Seychelles.
(10):279-393. Leiden, Netherlands: 65-74.
HOOPER,J.N. A. 1996.A toxic BiemnafromMadagascar SOEST, R.W.M. VAN & N. STENTOFT, N. 1988.
(Demospongiae: Poecilosclerida). In: Ph. Willenz (ed.). Barbados Deep-Water Sponges. Stud. Fauna Curasao
Recent advances in sponge biodiversity inventory and Carib.Isl. 70(215): 1-175.
documentation.Bull. Inst.r. Sci. nat. Belg., Biologic 66 THOMAS, P.A. 1973. Marine Demospongiae of Mahe
(suppl.): 123-133. Islandin theSeychelles Bank(IndianOcean).Annls. Mus.
KELLER, C. 1889. Die Spongienfauna desrothen Meeres r. Afr. cent.,Sci. zool. 203: 1-96.
(I.Halfte).Z.wiss. Zool. 48: 311-405. TOPSENT, E. 1897. Spongiaires de la Baie d'Amboine.
LEVI, C. 1963. Spongiaires d'Afrique du Sud. (1) (Voyage de MM. M. Bedot etC. Pictetdans l'Archipel
Poecilosclerides. Trans.R. Soc. S.Afr. 37(1): 1-72. Malais.) Rev. suisse Zool. 4: 421-487.
LEVI, C.& P.LEVI, 1989. Spongiaires (MUSORSTOM 1 VACELET. J. & P VASSEUR, 1971. Eponges des recifs
& 2). In: Forest, J. (ed.). Resultats des Campagnes coralliens de Tulear (Madagascar).Tethys (suppl. 1): 51-
MUSORSTOM,Volume 4.Mem. Mus. natn. Hist. nat. 126.
Paris (A, Zoologie) 143: 25-103.
Received: April 4, 2002
24