The Sponges of Micronesia, Part I The Palau Archipelago

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The Sponges of Micronesia, Part I
The Palau Archipelago
PATRICIAR.BERGQUIST!
ALTHOUGH potentially interesting from a zoo In order to generalize about the relation
geographic standpoint, the marine fauna of the ships of the Palau Islands sponge fauna it is
Palau Islands and of the islands and atolls to necessary to know something of the faunas of
the eastward, with the exception of Hawaii, contiguous areas. Two main regions have con
have received scant attention from expeditions tributed to the Palauan fauna, the Indo-Malayan
and individual collectors. It was to repair this and the New Guinea-northern Australian. The
gap in our knowledge that the "Project Coral sponge fauna of the Indo-Malayan region is
Fish" expeditions to the western and central known chieflyfrom the works of Thiele (1898,
Pacific were undertaken. The first of these ex 1900, 1903), Lindgren (1897), Kieschnick
peditions visited the Palau Islands in 1955 and (1898), Topsenr (1897), Hentschel (1912),
made extensive marine collections, of which Wilson (1925), de Laubenfels (1935), and
the sponges described below were a part. Levi (1%1).
Previous work relating to the Demospongiae Islands to the east of the Palaus are rela
of the Palau Islands is restricted to de Lauben tively unknown in respect to their sponge
fels' (1954) "Sponges of the West Central fauna. With the exception of Hawaii the lit
Pacific." This monograph was based upon the erature is restricted to Kirkpatrick (1900) for
author's own collections in the Caroline Islands, Christmas Island; and de Laubenfels (1949)
Marshall Islands, and the Marianas; and it con for Yap, (1955) for Onoroa, and (1954) for
tains, in addition to descriptions of many new the Marshall Islands, Caroline Islands, and the
species, significant field observations which Marianas.
have facilitated identification of the' present Our knowledge'"of the New Guinea and
collection. There are 33 species common to the northern Australian fauna stems from the work
two collections and the identification of these of Ridley (1884) and Burton (1934).
has necessitated considerable revision of por Many of the references cited above are very
tions of de Laubenfels' monograph.
old and require extensive revision; this, added
The Palau Islands belong to the Western to the obviousincompletenessof the faunalpic
Caroline group and aresituated on a submarine
ture they present of their respective areas, re
ridge running northeast from the Moluccas at
stricts the zoogeographic comparisons that can
6° 53' N to 8° 06' N,'134° 29' E. Between
be made. In view of the unreliability of the
the Palaus and the Philippines is the Philip
literature, the statements made here with re
pine Trench, which acts as an effective natural
barrier to the free migration of shallow-water spect to the Palau Islands are based solely on
invertebrates. Consequently, zoogeographic in the specimens examined in the course of this
investigation, and draw on de Laubenfels' work
terest in the Palau area focuses on whether the
only where this has been verified. It will be
fauna has remained similar to that of the Indo
possible to give a more detailed analysis when
Malayan region, through interchange of popu
lations across the intervening 600 miles, or has collections from Haluk, Kapingarnarangi, the
Philippines,and Hawaii havebeen fullystudied.
diverged significantly, producing a high per
centage of endemic species. Fifty species are described in this report; of
this number nine are new.A total of 14 species
Zoology Department, University of Auckland,
1 is endemic to the Palau Islands; seven species
New Zealand, and Division of Invertebrate Zoology,
are known only from the Palaus eastward, in
PeabodyMuseum ofNatural History,YaleUniversity.
Manuscript receivedSeptember 6, 1963. the case of Xestospongia exigua eastward to
123
124 PACIFIC SCIENCE, Vol. XIX, April 1965
Christmas Island. Thirteen species are common STATION DATA
to the Indo-Malayan and Palau areas, extend
Sta,10. Madalai district, west end of Koror 1.,
ing, in the case of lotrocbota ba::ttlife.ya, Clatb
Abe's Traverse III.Sand bottom with Enhalus
ria fasciculata,Clatbria cervicornis,Neofolitispa
and Halimeda; depth 12 inches at low water.
.diancbora,and Asteropus sarasinorum, eastward
July 8, 1955.
to the Marshall Islands. Eleven species have a
Sra,12. Madalai district, west end of Koror 1.,
wide Indo-Pacific distribution: Heteronema
Abe's Traverse II, at south end of Koror
erecta, lrcinia ramosa, Phyllospongia foliascens,
Arakabesan Causeway. Mud and sand, with
Psammaplysilla purpurea, Dysidea berbacea,
Enhalus, Halimeda and Padina; exposed at
Coelocarteria singaporense, Myrmekioderma
low water. July 9, 1955.
granulata, Spirastrella vagabU1uia, Placospongia
Sta.15. Madalai district, west end of Koror 1.
melobesioides, Ancorina acervus,and Cinachyra
Shallow flat enclosed by retaining wall par
australiensis.Threespecies are known elsewhere
allel to Malakal Causeway. Sand and silt,
only from Australia, in the case of [as-pis coria
with Enhalus; depth 18-24 inches at low
ceaonly from South Australia.
water. July 11, 1955.
The sponge fauna of the Palau Islands is
Sta.25. Inner margin of barrier reef, 8 miles
certainly richer and more varied than that of
NW of Koror. Sand, coral,dead coral blocks;
any of the more easterly islands thus far stud
depth 3-7 ft. July 19, 1955.
ied. It is not possible, however, to compare the
Sta.35. Peleliu boat channel, between Ngarger
quantity and variety of Demospongiae in Indo
sal and Kongauru islands, ca. 1 mile east of
Malayan localities with that of the Palau Is
north tip of Peleliu. Sand, silt, some coral;
lands without more data on both areas.
Enhalus acoroides and 2 other species of eel
The broad system of classification used in
grass; algae incl. Caulerpa spp. Depth 1-10
this work is basically that propounded by Top
ft. July 24, 1955.
.sent (1928) and modified by de Laubenfels
Sta.47. Iwayama Bay, shore of islands XXXIII
(1936). In several cases minor alterations in
and XXXIV,west side ofKogai-Hamo.Sand,
this system have been suggested. The Axinelli
live and dead coral; Enhalus in sand, Padina
.dae are still treated as a family of the Hali
on rocky areas.Tuly 28, 1955.
chondrida, although it is clear that they should
Sta.53. Ngaremdiu area, eastern Urukthapel;
be elevated to the level of a separate order. To
rocky cape west of sand beach Oirarel-ruul.
reassign the genera involved is a long task, the
Sand, coral, rock; small eel grass and Halo
material for which is not immediatelyavailable.
phila ovalis; depth 0-7 ft. July 31, 1955.
Throughout the text means are given in pa
Sta. 59. Reef bordering eel grass flat east of
rentheses with the range of spicule dimensions.
Ebadul's dock, north shore of Koror. Living
All color notations are after Munsell's Book
and dead coral, with sparse patches of sand.
of Color.
Aug.4, 1955.
Sea. 50. Sand and eel grassflat west of Ebadul's
ACKNOWLEDGMENTS
dock, north shore of Koror. Sand with occa
The author is greatly indebted to Dr. W. D. sional coral patches, Enhalus abundant, long
Hartman for his advice and assistance through streamersofsargassum attached to dead coral.
out the preparation of this manuscript. August 5,1955.
Mr. R. Binkowski is thanked for technical Sta.61. Seaward reef flat at south end of Nge
assistance, Mr. J. Howard for photographic melis 1., west side of Palau Archipelago.
work, and Mrs. Shirley Hartman for the prep Coral and sand, some Enhalus and Halimeda
aration of all drawings. in sand patches; depth 18 inches-6 ft.August
Thisresearch wasmade possiblebytheaward 6, 1955.
of a Sessel Fellowship in Biology from Yale Sta,64. Small bay at southern end of Eil Malk
University and by National Science Foundation lagoon. Bottom limestone and sand, little
Gram GB-516 to Yale University. coral; depth 6-20 ft. August 7, 1955.
Sponges of Palau,i-BERGQUIST 125
Sta.67. Reef in pass west of Nghus ("Ankosu" rocksof foreign origin; depth 4-5 ft.August
on charts), southern tip of Urukrhapel. Liv 30, 1955.
ing and dead coral, sand, coral rubble; depth Sta.219. Iwayama Bay (Abe's Division L), ex
5-7 ft. Aug.7,1955. treme eastern part.Marginal reef along coast
Sta.92. Iwayama Bay,south end of island XV; of Koror called U'larnii' (or Uchlamiich).
Abe's Traverse XIII. Bottom silty sand over Living and dead coral, sand patches; depth
rocky limestone platform, marginal living 3-20 ft.October 12, 1955.
reef; Enhalus, Padina, Halimeda; depth 3-20 Sta.220. Iwayama Bay, east side of mouth of
ft.August 14,1955. Kaki-suido (Oyster Pass) between islands
Sta.100. Iwayama Bay,Bay of the Dragon Pal XXIX and SE Koror. Limestone, live and
ace, west side of Kogai-hanro. Bottom coral, dead coral; depth 3-20 ft. October 12, 1955.
sand in pockets; depth 5-15 ft. August 16, Sta.220A. Same as 220 but collected OCt. 22,
1955. 1955.
Sta.104. Malakal Harbor.From sunken ship off Sta.220B. Same as 220 but collected Oct. 29,
north shore of Urukthapel; depth 40-50 ft. 1955.
August 17,1955. Sta.236A. Same as 220 but 150 yds. north.
Sta.106. Reef flat, Ngadarak reef north of Oct.20, 1955.
mouth of Malakal Pass. Exposed coral rubble Sta.245. West shore of Urukthapel, in small
with residual tide pools. August 17, 1955. bay; 7° 16' 31/1 N, 134° 26' 13/1 E. Lime
Sta,111. Reef flat, Ngaremdiu reef,east side of stone shelf,coral, Halimeda, 3-10 ft. October
Urukthapel.Exposed boulder flat. August 19, 23,1955.
1955. Sta.252. Fringing reef of small island in west
Sta.124. Yoo Passage, west of KasaoReef, 3Y4 part of lagoon of Eil Malk Island.Limestone,
miles SSE of Ngaremdiu, east side of Uruk coral, sand,0-10 ft.October 27, 1955.
thapel. August 24, 1955 [Gloria Maris Sra. Sta.258. Small bay in north coast of Ngarem
450, haul 2]. diu region of Urukthapel, near Japanese
Sta.125. 1% miles NE of Ngabadangel (Cape stone pier. Sand, live and dead coral, lime
"Gabadagur.n" on charts). Sand with Hali stone, with Enhalus and Halimeda, much
meda and S~;'iatopora, 17 fms. August 24, stinging Stephanoscyphus; depth 2-4 ft.No
1955. [Gloria Maris Sta. 452, haul 1). vember2,1955.
Sta.133. Iwayama Bay,south shore of island II.
SYSTEMATICDESCRIPTIONANDDISCUSSION
Reef flat between shore and deep reef pool;
(Species includedarelisted in systematic order)
area of Abe's traverses VIII, IX, and X.
Coral with sand pockets; depth 2-3 fr at OrderKeratosa Grant
m.l.w.August 28, 1955. SuborderDictyoceratidaMinchin
Sta.134. Iwayama Bay,south end of island XX. FamilySpongiidaeGray
Reef shelf and slope; bottom entirely coral Genus SpongiaLinne
and rock; depth 3-20 ft.August 28, 1955. Spongia officinalisLinne
Sta.135. IwayamaBay,westsideofislandXXII. Genus Dactylospongia n.gen.
Reef shelf and slope; bottom entirely coral Dactylospongiaelegans (Thiele)
and rock; depth 3-40 ft. August 28, 1955. Genus HeteronemaKeller
Sta.136. Reef flat outside of large cave at SE HeteronemaerectaKeller
end of Koror, east entrance of Iwayama Bay. Genus IrciniaNardo
Sand, branched coral (Montipora, Seriato IrciniaramosaKeller
pora), and vegetation (Enhalus, Halimeda, GenusPhyllospongia
Padina);depth 4-6 ft.August 28,1955. Phyllospongiafoliascens (Pallas)
Sta.140. Iwayama Bay,in Ngerchelngae1 ("Ge PhyllospongiadendyiLendenfeld
ruherugairu" on charts) Pass,between Kogai Genus Fasciospongia Burton
Peninsula and Kaibakku I. Montipora-flat Fasciospongiacbondrodes
with Enhalus, Halimeda, Padina, and some (deLaubenfels)
126 PAOFIC SOENCE, Vol. XIX, April 1965
Genus PsammaplysillaKeller SuborderMicrocioniformes de Laubenfels
PsammaplysillapurpureaCarter FamilyMicrocionidaeHentschel
FamilyDysideidaeGray GenusMicrocionaBowerbank
Genus DysideaJohnston Microcionaeurypa (deLaubenfels)
Dysideaberbacea (Keller) GenusClathria Schmidt
DysideachloreadeLaubenfels Clathriacervicornis (Thiele)
Dysideagranulosan.sp. ClathriafasciculataWilson
Dysideaarenarian.sp. Family OphlitaspongiidaedeLaubenfels
GenusEuryspongia GenusMycaleGray
Euryspongialobatan.sp. Mycalelissochelan.sp.
Mycalecavernosan.sp.
Order Haplosc1eridaTopsent
GenusNeofolitispa nom. nov.
Family HaliclonidaedeLaubenfels
Neofolitispadianchora
GenusHaliclonaGrant
(deLaubenfels)
Haliclonavetinea,(de Laubenfels)
GenusDesmacellaSchmidt
HaliclonakoreinelladeLaubenfels
Desmacellalampra deLaubenfels
Genus Cribrochalina Schmidt
CibrochtilinaolemdadeLaubenfels OrderHalichondridaTopsent
Genus XestospongiadeLaubenfels Family AxinellidaeRidley and Dendy
Xestospongiaexigua (Kirkpatrick) GenusPseudaxyinissaBurton
Family Callyspongiidae deLaubenfels PseudaxyinissapitysdeLaubenfels
GenusCallyspongiaDuchassaingand Genus PhycopsisCarter
Michelotti Phycopsissp.d.terpnis de Laubenfels
Callyspongiasubarmigera (Ridley) FamilyDesmoxyidae Hallmann
CallyspongiaridleyiBurton GenusHigginsiaHiggin
FamilyDesmacidonidaeGray Higginsiamixta (Hentschel)
GenusGelliodesRidley Genus MyrmekiodermaEhlers
GelliodesgracilisHentschel Myrmekiodermagranulata (Esper)
OrderPoecilosc1eridaTopsent OrderHadromeridaTopsent
Suborder PhorbasiformesdeLaubenfels FamilySpirasrrellidae Hentschel
FamilyAdoeiidae deLaubenfels GenusSpirastrellaSchmidt
Genus AdociaGray SpirastrellaaurivilliLindgren
Adociaturquosiade Laubenfels SpirastrellavagabundaRidley
Genus ToxadociadeLaubenfels GenusTimeaGray
ToxadociaviolaceadeLaubenfels Timeagranulatan.sp.
Genus Orina Gray Family PlacospongiidaeGray
OrinasagittariaSollas Genus PlacospongiaGray
Genus Kallypilidionde Laubenfels PlacospongiamelobesioidesGray
Kallypilidion poseidon deLaubenfels
OrderEpipolasida Sollas
Genus PellinaSchmidt
Family JaspidaedeLaubenfels
Pellinacarbonaria (Lamarck)
Genus AsteropusSollas
Genus Siphonodictyon n.gen.
Asteropussarasinorum (Thiele)
Siphonodictyon mucosan.sp.
GenusJaspisGray
Family CoelosphaeridaeHentschel
Jaspiscoriacea (Carter)
GenusCoelocarteriaBurton
Coelocarteriasingaporense (Carter) OrderChoristida Sollas
SuborderMyxilliformesdeLaubenfels FamilyAncorinidaeGray
FamilyTedaniidaeRidleyand Dendy Subfamily AncorininaedeLaubenfels
Genus lotrochotaRidley GenusAncorinaSchmidt
lotrochota baculiferaRidley Ancorinaacervus (Bowerbank)
Sponges ofPalau, I-BERGQUIST 127
SubfamilyStelletrinaeSallas GENUS Dactylospongianov.gen.
GenusStelletta Schmidt
Dictyoceratida with skeleton a relatively reg
Stellettadurissima n.sp.
ular network of polygonal meshes without dif
Family TetillidaeSallas
ferentiation into ascending and connecting ele
Genus TetillaSchmidt
ments. Fibres lack foreign inclusions except for
Tetillamicroxean.sp.
a few isolated spicule fragments, have a granu
Genus ParatetillaDendy
late surface textureand a faintly stratified struc
Paratetilla bacca(Selenka)
ture. The sponge surface is free of detritus and
Genus CinachyraSallas
is covered with irregular conules. The dermal
Cinachyraaustraliensis (Carter)
membrane between rows of conules is stretched
(complexofspecies)
over deep subdermal channels which extend for
aconsiderableverticaldistance. Flagellate cham
ORDERKERATOSA Grant bers are evenly distributed throughout the endo
SUBORDERDlCTYOCERATIDAMinchin some, and are spherical and small, 24-30,... in
FAMILYSPONGIIDAEGray diameter. The genus is erected for the type spe
GENUSSpongia Linne cies, Luffariella elegans Thiele, and is most
closelyallied to Hippospongia.
Spongia officinalis Linne, 1759, p. 1348,pI.1,
Dactylospongiaelegans (Thiele)
RESTRICTED,SYNONYMY:
Spongia~ffi~inalis Fig. la, b
Linne,1759, p.1348,pI. 1,
figs.1and2. Luffariella elegans Thiele, 1899, p. 25, pI. 3,
Spongia officinalis subspecies matamata de fig.4,pI.5,fig.20.
Laubenfels, 1954,p.4.
OCCURRENCE:Sta.47.Palau Islands.
DESCRIPTION: Several pieces of this sponge
OCCURRENCE: Sta.60.
were obtained, probably all from one large col
REMARKS: A single specimen of this sponge
ony with long thin anastomosing branches di
is in the present collection. It compares well
verging from a small poorly defined basal
with the type of de Laubenfels' S. officinalis
region. The branches are 0.5-1.5 cm wide, up
subspecies matamata, USNM 23200, except that
to 32 cm long, and many have several anasto
the primary fibres are stouter, up to 80,... in mosesalong theirlength.
diameter, and more frequent than is inferred COLOR: In alcohol, dark reddish-brown, be
in de Laubenfels' (1954) description. tween (rY-R2/2) and (rY-3/2).
Embryos in all stages of segmentation are TEXTURE: Extremely tough and elastic, al
present in the specimen, and the older embryos most rubbery.
are verydarkly pigmented. SURFACE: Coarselyconulose,with the conules
No attempt ismade here to evaluate the pre tending to be aligned in rows between which
vious records of Spongia officinalis from the the dermal membrane is stretched. In parts of
Indo-Pacific region. It isevident that systematic the sponge the dermal membrane of either sur
face is all that connects two adjacent branches.
relationships are confused in the entire genus
In the preserved sponge the membrane is
and particularly in S. officinalis and S. zimocca.
sunken into deep subdermal cavities which may
A careful study of more extensive collections
be up to 4.0 ern long and 0.5 cm wide. Conules
than are presently available is necessary before
are irregular, squarish, multituberculate, and
affirming that all specimens referred to these
up to 0.5 ern long, 0.2 ern high, and 0.2 em
species are correctly assigned.
wide. The oscules are distributed irregularly
DISTRIBUTION: Mediterranean, West Indies, over the surface of the branches and are 0.08
Australia, Indian Ocean, Norway(?), South 0.2 mm in diameter. Pores are absent from the
America, Marshall Islands. areas where the dermal membrane extends over
128 PAOFIC SOENCE, Vol. XIX, April 1965
Considerable confusion has centered around
the correct generic position of Luffaria vari
abilis. The genus Luffaria was unrecognizably
describedbyDuchassaing andMichelotti(1864).
Schmidt (1870) redeseribed the genus with
Out adding any species, and Polejaeff (1884)
described Lttffaria variabilis. Lendenfeld (1889)
recognized that Polejaeff's sponge was not a
Luffaria as understood by Schmidt (= Veron
gia), and took the erroneous step of re-estab
lishing Luffaria under Polejaeff's name to reo
ceiveL. variabilis and tWO new species.Topsent
(1934) considered that both Luffaria and Luf
fariella should be abandoned and all species in
volved transferred to Aplysinopsis Lendenfeld.
The difference between Aplysinopsis and Tho
recta Lendenfeld is very slight, merely that
Thorecta has a relatively smooth surface and
more foreign material in the dermis (see de
Laubenfels, 1948). These two genera are now
consideredsynonymous.
The type specimen of Luffaria variabilis
Polejaeffshould be established as BM 85.8.8.52.
This sponge has been examined and in most
FIG. la. Dactylospongia elegans (Thiele). USNM
features it is compatible with Cacospongia. It
23707.
has, however, one distinguishing characteristic:
subdermal cavities, bur"~bundant over most of
an extremely fine and regular tertiary fibre net
thesurface.
work is present throughout the sponge. In
SKELETON: A compact irregular reticulation view of this marked difference in the skele
of clear yellow-brown fibres; many ascending
ton, it is proposed to retain Luffariella for L.
fibres are distinguishable only immediately be
variabilis and other sponges which have com
low the surface.Some stratification is shown in
parable skeletal structure. Lufjariella geometries
most fibresand a poorlydemarcated pith occurs
Kirkpatrickmayproperlybelonghere.
infrequently. Isolated spicule fragments occur
It is clear that Luffariella elegans Thiele is
randomly. A characteristic feature of the skele
generically distinct from Luffariella variabilis.
ton is the granulate surface of the fibres. Fibre
Examination of the Palau specimen of L.
diameter is 20-54ft (38ft).
elegans and an undescribed specimen in the
FLAGELLATE CHAMBERS: Small, spherical,
British Museum (1946.11.25.170) shows that
24-30ft in diameter, and are regularly distrib
the correct generic grouping for this sponge is
uted throughout the body of the sponge except
not near Cacospongia but near Hippospongia,
in the lacunar areas which surround the sub
differing from some specimens of this genus
dermal cavities. The endosome contains many
only in form, the presence of long surface chan
darkly pigmented granules and abundant dia
nels, and the lack of any cored primary fibres.
tomskeletons.
These are the only features separating L. ele
DISCUSSION: Thiele (1899) described this
gans and Hippospongiametachromies (de Lau
sponge from the Celebes and established the
benfels), also from the Palau Islands.
genus Luffariella to receive Luffaria variabilis
Polejaeffand Luffariella elegans; the formerwas
No cored ascending fibres were described for this
2
designated as the type of the genus by Thiele
sponge, but re-examination of the type has shown
and later byde Laubenfels (1936). them to be present in the subdermal region.
Sponges of Palau,I-BERGQUIST 129
Since the genus Luffariella is retained for L. are small and spherical, 20-30,.. in diameter;
variabilis,a newgenusDactylospongia iserected the subdermal region is extremely cavernous:
to receive Lufjariellaelegans. and darkly pigmented. This sponge is not easily
DISTRIBUTION: Celebes (Thiele) distinguishable from Heteronema. '
The second species described in Tborecto
GENUSHeteronema Keller
psamma is T. chromogenia de Laubenfels from
Heteronema erectaKeller Bermuda. This sponge is a dark red color and
Fig.2 has far less debris in the fibres than T. irregu
Heteronema erectaKeller, 1889,p.339. laris.The flagellate chambers were undescribed
Heteronema erectaRow, 1911,p.369. and apparently much sand was incorporated in
Thorectopsamma mela de Laubenfels, 1954, the body of the sponge. It is unlikely that this
p.29,fig.15,pI.8,fig.6. sponge belongs toThorectopsammabut, in view
Thorectopsamma mela de Laubenfels, 1955, of our incomplete knowledge of its anatomy,
p.138. no suggestion can be made about its real affin
ities. Thorectopsamma xana de Laubenfels:
OCCURRENCE: Sta.10,100,140. (1954) is referred to Psammaplysilla purpurea.
DESCRIPTION: De Laubenfels (1954) has DISTRIBUTION: Red Sea (Keller, Topsenr,
given an excellent description of this sponge Row); West Central Pacific; Hawaii (de Lau
as it occurs in the Micronesian area and little benfels).
can be added except to note that: (1) the pri GENUSlrcinia Nardo
mary fibres have a tendency to be fasciculated
lr iniaramosa (Keller)
immediately below the surface; (2) The dermis
is crowded with granules containing a black
pigment; (3) The immediate subdermal region
is cavernous, consequently the dermis is easily
detachable; and (4) The pigment granules in
the subdermal and deep layers of the body are
extremely abundant and tend to obscure the
chambers and canalsystem.
DISCUSSION: Comparison of Red Sea speci
mens of Heteronema erecta with de Lauben
fels' Pacific sponges identified as Tborecto
psamma mela reveals that the two species are
identical. The extremely characteristic surface
pattern of radiating ridges between conules is
present in all specimens examined from both
geographical areas. The texture of the sponge
varies greatly: some of the Palau specimens
are relatively compressible; others are extremely
hard and thus compare with specimens de
scribed by Row (1911).
With the removal of T.mela to Heteronema,
the genus Thorectopsamma Burton is reduced
to three species.T. irregularis,the type species,
was described from a single specimen; it had
laminated fibres densely packed with debris and
no special dermal skeleton. Burton made no
mention of the soft tissues of his sponge. Re
examination of the type of T. irregularis (BM FIG. lb. Dactylospongia elegans (Thiele). Portion
30.8.13.217) shows that the flagellate chambers of the fibre network.
130 PAOFIC SOENCE, Vol. XIX, April 1965
RESTRICTEDSYNONYMY: benfels', or Keller's specimens; the flagellate
Hircinia ramasaKeller, 1889, p. 345, pI. 20, chambers range from 24-36fL in diameter.
fig.5. De Laubenfels (1950) differentiates 1. ra
IrciniaramasadeLaubenfels, 1948,p. 73. mosa,in the West Indies, from 1.fasciculata by
Irciniaramosade Laubenfels,1954,p.23,fig. the form of the branch tips, rounded in ramosa,
11. pointed in fasciculata. Hartman (1959) has
emphasized the presence of a dermal reticulum
OCCURRENCE: Sra,15,60,92.
of sand grains in I.ramosa. At present it is im
REMARKS: De Laubenfels (1954) gives a
possible to be sure that the Pacificsponges dis
relatively detailed description of this species
cussedabove belong to the samespeciesasWest
and stresses the ramose form, the abundant fine
Indian sponges also named I. ramasa; the same
filaments, and the presence of a second coarser
type of filament containing refractile material problem applies to I. fasciculata. In the case of
as the characteristic features. The specimens in ramosa, however, the Pacific sponges seem
this collection agree in most details with the closertoKeller's originaldescription and, in the
above.Thefibreslacking granules haveagreater event some specific difference is found, the
range in diameter, 2.0-5.0fL, than either deLau- West Indian population would require a new
FIG.2. Heteronema erectaKeller. Left, Sta. 10;right,Sta. 140.
SpongesofPalau,I-BERGQUIST 131
FIG.3a. Phyllospongi«[oliascens (Pallas).Sta. 111.
FIG. 3b. Pbyllospongi« [oliascens (Pallas). Upper,
Sta. 245; lower, Sta. 220A.
specificname. It is difficult to see what charac
ters can be used to distinguish between such
sponges as I. ramosa, 1.fasciculata, and I. den DESCRIPTION: Three of the five specimens
droides since the entire genus is extremely uni are similar in form to P;foliascens asillustrated
form in internal morphology and, seemingly, by Lendenfeld (l889:pl. 5, fig. 3), one is
variable in habit and surface characteristics. almost identical to P. lekanis de Laubenfels
Studies of the ecology and morphology of liv (l954:pl. Ill, fig.a), and one, of spiral shape,
ing populations are urgently required in this answers to de Laubenfels' description of large
genus. specimens of P. lekanis which he observed in
DISTRIBUTION: Red Sea, West Indies, Palau the field. There is a great variation in the sur
Islands,Ponape,GreatBarrier Reef. face ornamentation, and in the distribution and
abundanceofoscules.In allspecimens theskele
GENUSPbyllospongi«Ehlers
ton is compact, with cored primary tracts and
Phyllospongia foliascens (Pallas) thick sand cortex on both surfaces. In this re
Fig.3a, b spect the present specimens contrast with the
holotype of P. lekanis (USNM 23109), which
RESTRICTEDSYNONYMY:
has a loose skeleton mesh, lightly cored fibres,
Spongia[oliascens Pallas,1766,p.395.
and a thin sand cortex on both surfaces.
Phyllospongia foliascens Lendenfeld, 1889, p.
Little note has been taken in the systematics
196, pl. v, fig. 3; pI. vi, figs. 1, 3, 4, 10; pI. vii,
of Phyllospongia of the amount of debris pres
fig. 11;pI.xiv,fig.2;pl.xxiv,fig.6.
ent in the cortex and the skeleton; this, in con
Carteriospongia foliascens Burton, 1934, p.
junction with the range of form exhibited by
573.
this sponge in the Palaus, is sufficient reason
Phyllospongia lekanis de Laubenfels, 1954,p.
for relegating P.lekanis to P. foliascens..
15,fig.7,pl. 111,fig.a.
DISCUSSION: It is with some hesitation that
OCCURRENCE:Sta. Ill,220A, 245. these specimens are described as Phyllospongia
132 PAOFIC SCIENCE, Vol. XIX, April 1965
rather than Carteriospongia.Itseems that Spon
gia foliascens Pallas, which is the type species
of Carteriospongia Hyatt by subsequent desig
nation of Burton (1954),hasabundant foreign
material incorporated into the dermis and the
primary fibres. Spongia papyracea Esper, the
type species of Phyllospongia Ehlers, is under
stood by Burton to have no sandy inclusions in
either the fibres or the cortex. Ehlers (1870)
does not comment specifically on this feature.
Many authors (Dendy, Ridley, Bowerbank, de
Laubenfels) have described sponges with cored
ascending fibres as P. papyracea. De Laubenfels
(1948) suggests that the type specimen is one
in which primary fibres are rare or absent and
that the specimens described by later authors
are congeneric with Esper'ssponge even though
they possess varying amounts of debris in both
cortex and primary fibres. The commonly ac
cepted view of Phyllospongia is of a sponge
with a variable amount of debris in both fibres
and cortex, and in keeping with this view the
Palau specimens are assigned to Phyllospongia.
A restudy of Spongia papyracea Esper is re
quired to clarify this problem.
DISTRIBUTION: Red Sea, Indian Ocean, Ma
laya, Australia, New Zealand," West Central
Pacific. FIG. 4a. Lower, Pbyllospongia dendyi Lendenfeld.
Sra, 92. Upper, Fasciospongia cbondrodes (de lau
Phyllospongiadendyi Lendenfeld benfels).Sta.92A::.
Fig.4a,b
ing, particularly the outer series. The single
Phyllospongia dendyi Lendenfeld, 1889, p. specimen is 10 cm long, 5.0 em wide and 6.0
177,pl.14,fig.5. em high; lamellae are 2.0-3.2 mm thick. .
Phyllospongia dendyi var. frondosa Lenden COLOR: In alcohol, dark reddish-brown, be
feld,1889,p.178,pl.14,fig.5. tween (rY-R 5/2) and (rY-R5/3).
Phyllospongia dendyi var. digitata Lenden- TEXTURE: Fleshy but sufficiently elastic for
feld, 1889;p. 178. the sponge to regain its shape after being
Phyllospongia dendyi Burton, 1934, p. 573. compressed.
OCCURRENCE: Sra,92. SURFACE: Covered evenly with sharp fine
conules up to 0.5 mm high. Oscules are scar
DESCRIPTION: This sponge consists of some
tered over the sides of the lamellae and range
what inclined lamellaedecreasing in heightfrom
from 0.2-1.2 mm in diameter. Narrow sub
the center to the margin and often bearing lobe
dermal channels are visible through the dermal
like secondary lamellae on the outer surface.
membrane; these meander over the surface in
The lamellae arise from a spreading base and
certain areas and are absent near the edges of
tend to intersect to form a series of pouches.
thelamellae. .
The free margins of the lamellae are undular-
SKELETON: Composed of primary fibres
cored with spicule fragments and running diag
8This record isdoubtful; Lendenfeld described sev
eral species and varieties of Phyllospongia from New onally through the lamellae.Theyare connected
Zealand, none of which has been verified. by a network of uncored secondary fibres.