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MUS. Comp. ZOOL:
LIBRARY

AUG 3 1 1973

HARVARD
UNIVERSITy

POSTILLA

PEABODY MUSEUM
YALE UNIVERSITY

NUMBER 161 29 AUGUST 1973

THE SYSTEMATICS OF SOME
JAMAICAN EXCAVATING SPONGES
(PORIFERA)

ROSEMARY K. PANG

POSTILLA

Published by the Peabody Museum of Natural History, Yale University

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THE SYSTEMATICS OF SOME JAMAICAN
EXCAVATING SPONGES (PORIFERA)

ROSEMARY K. PANG*

Department of Biology, Yale University, New Haven, Connecticut

(Received 21 January 1972)

ABSTRACT

Thirteen species of coral-dwelling sponges are reported from the reefs of
Discovery Bay on the north coast of Jamaica. Seven of these are new species:
Cliona delitrix, C. peponaca, C. langae, C. laticavicola, C. aprica (family Cli-
onidae); Alectona jamaicensis (family uncertain); and Siphonodictyon brevitubu-
latum (family Adociidae). Two other species, Cliona schmidti and C. janitrix, are
new records for the West Indies.

A shallow-water and a deep-water form of two new species are described.
Transplant studies indicate that these forms, distinguished by differences in the
dimensions of the spicules and papillae, the degree of papillary fusion and, to a
slight degree, spicule morphology, are largely habitat-induced. It is suggested
that differences in silica concentration with depth may affect spicule sizes.

*Present address: Dept. of Pharmacology, College of Physicians and Surgeons
of Columbia University, New York, N.Y. 10032.

POSTILLA 161: 75 p. 29 AUGUST 1973

Contents

To Introduction x. dicccgeSsccecacscnaesscutce vesecseteeeetnncsnuehassetece auentsaaeee eee 3
Il. Previous records of excavating sponges from the West Indies, Florida

and the Gulf Of MEXICO siccc.c.aceesceesensedieecansnsenscevabe nassseetel eee eee eee 5)

III. Descriptions of Jamaican species of excavating SPONQES ............:00ceeeeee 8

A. Cliona schmidti (Ridley) Topsent «....25.0-:.015-.2s0ccom.soeeeee ee eee 8

B: Cliona vermifera Hancock. 22.0.0 .ssescsss.o1.caeseossse0e leek cae eee eee 12

C. -Cliona janitrix Topsent wiac.cesistec.scedsesssooeseses ater dectonneiea ests eae een 16

D. GCliona lampa de Vaubeniels. s:....5...ccs0cs-necssoceassincstnfenseteneseeeeseeeeeteeeamet 18

E.’ Cliona caribbaed Carters. ve.sc. sicesceccess02500s008 <s -00vedenonass see arn eee ee eeeeee ap)

F) Cliona@’ Gelttrix, 1.8 Ps sicc.cisasecteee-coccesecudeasanseseasees ones) sadece eee mee aeeeaeeene 28

GG, Cliona Pemonacd, DSP s- sacecacze.sqa's3s%a.50heesesesdae<38s5-2-asaeap see eee eee 32

Hi... Cliona lang@é, WSs 2.1 s.s.dsccccecssemescdesaas sncesisveessaeseoiesonesteaene pete eee 34

Il. Cliona laticavicola, n:Sp. i.ic.cesssisess icone. ceeusesnns ooene esse ce eee eee 37

J. CHONG ApricG, MSP. x... cc2c<edeasce.s2aclsosobsscssvesseers sce «28 snc0e ceeseet eee eeneeneeee 42

K. Anthosigmella varians (Duch. & Mich.) de Laubenfels .................... 47

L.. Allectona jaMaiCensis, DSP. ....22..0+sssec0e0+s095¢9shdnassseunasdesaa eee eee 50

M. Siphonodictyon brevitubulatum, NSP. ..:00....-50005 s000 ++ os suse reua eee 56

IV.. Key to species described | .s........cscsncdeiesssne (Ssoncovcbousass secede seesslaseetteeeaeeeteeteneet 58

V. Diversity and ZOO gEOSTA PHY: wicc.svecwassssossneseunaecoteemncnoshacntesnsee eeeeteee ene 60

VI. Transplant StudieS. ..2:......:c025:.0<csesecccans soboseastunensavessdecosessea ace ehenenetatepeeeeee 65

VIN. Acknowledgements, oc. ucsc.c.ceeicsenesennseeedee'cyaeeoueser sends cecssaeece eteeeeeeteeeeeeenenen 68

VIM. Literature Cited 2.3.0.5... cseenas eben decneine'sudecccdob ts .nds Ade oncsnte ce tecew ent aaees a eneee ence 68

JAMAICAN EXCAVATING SPONGES 3

I. Introduction

Excavating sponges of the family Clionidae (Subclass Tetractinomorpha, Order
Hadromerida, Lévi, 1957) have long been known to exert an important influ-
ence on the dissolution and recycling of accumulated calcium carbonate,
particularly that from oyster beds and coral reefs. Osler (1826) was the first to
recognize the role played by sponges in the destruction of oyster shells. His
report appeared only a few months before the genus Cliona was established by
Grant (1826) for the “new zoophyte”, Cliona celata, from the oyster beds of the
Firth of Forth, Scotland.

Hancock (1849) affirmed that the “new zoophyte”, Cliona celata Grant, was
indeed a sponge and not polypiferous as Grant had believed. He also stated that
C. celata was not an isolated, aberrant form but one of numerous species belong-
ing to the genus Cliona. Many of his species descriptions, based on dried speci-
mens gathered from the cabinets of British shell collectors, were inadequate.
Hancock’s type specimens were originally deposited in the Hancock Museum,
Newcastle-Upon- Tyne. However, when | visited the museum in August, 1969,
Dr. D. Grey, the zoologist, informed me that there were no records of these
specimens in the museum. Since Hancock’s clionid types are also not deposited
at the British Museum (Natural History), it may be assumed that they are lost.

The taxonomic history of the boring or excavating sponges has been both long
and complex. The voluminous literature reflects the many controversies and
debates that have taken place. More than two hundred species have been at-
tributed to Cliona and its synonymous genera (Vosmaer, 1933). Although the
establishment of the family Clionidae for the excavating sponges is usually
credited to Gray (1867), the name Clionidae was originally proposed by
d’Orbigny (1852) for sponges “qui percent l’intérieur des pierres et des coquilles
de canaux irreguliers ouverts de distance en distance par des oscules extérieurs”
(p. 209). The family contained only the genus Cliona. With the addition of
several new genera in subsequent years, the family Clionidae may now be divided
into two subgroups: (1) Cliona and related genera, and (2) Thoosa and related
genera, distinguished on the basis of differences in microscleres (de Laubenfels,
1936a).

The warm waters of the West Indian coral reefs with their abundance of cal-
cium carbonate substrates in the form of dead bases of living corals, dead corals,
calcareous algae, sclerosponges (Hartman and Goreau, 1970) and carbonate
rock, would seem to be an extremely favorable environment for the excavating
sponges as a group. However, previous investigations (de Laubenfels, 1936a, b,
1949, 1950a, b; Hechtel, 1965) have revealed only four species of clionids in this
region. Of these, one is endemic. De Laubenfels (1953a) and Little (1963), work-
ing in the Florida Gulf Coast region, reported two and five species of clionids,
respectively. Little’s fifth species, Cliona viridis (Schmidt) Gray, is probably a
suberitid (Hechtel, 1965).

By way of contrast, Volz (1939) reported nine (three endemic) species of
clionids from the shallow waters of the Adriatic sea, also a warm-water en-
vironment with considerable amounts of available carbonate substrate. Studies
of the temperate waters of eastern North America by Old (1941, 1942) and Hart-

4 POSTILLA 161

man (1958) revealed the presence of as many as four species of clionids inhabit-
ing the shells of oysters and other molluscs.

The systematics of the West Indian excavating sponges is largely unknown.
The present study enlarges our knowledge of the species occurring in one area of
the West Indian region, the reefs of Discovery Bay on the north coast of
Jamaica. Although the work of Goreau and Hartman (1963) indicated that
below a depth of IS meters, sponges are an exceedingly important constituent
of the north coast reef fauna of Jamaica, there are no previous records of
clionids from this area. During the course of the present investigation sponges
were collected by hand from depths of 0-50 meters using either free diving with
mask, snorkel and fins, or diving with SCUBA equipment. Thirteen species of
coral-dwelling sponges are reported; seven of these are new and are described
here. The geographical distribution and taxonomic history of the previously
described species are given.

Specimens are deposited in the Peabody Museum of Natural History, Yale
University, abbreviated as YPM in the text, the British Museum (Natural
History), abbreviated as BMNH, and the State University of New York-
University of the West Indies Marine Laboratory, Discovery Bay, Jamaica, W.1.,
abbreviated as SUNY-UWI.

Spicule measurements are reported as mean value with standard error,
followed by the absolute range.

JAMAICAN EXCAVATING SPONGES 5

II. Previous Records of Excavating Sponges from the West Indies,
Florida and the Gulf of Mexico

The shallow waters of the West Indian region possess a rich sponge fauna.
Although no previous studies have dealt exclusively with the excavating sponges
of this area, thirteen new species of clionids have been described from the West
Indies, Florida and the Gulf of Mexico by various investigators. Unfortunately,
ten of these species are insufficiently described. The list below summarizes all
the previous reports of excavating sponges from this region.

The earliest record of a West Indian clionid is that of Hancock (1849) who
described Cliona millepunctata from a single dried specimen found inhabiting a
West Indian helmet shell, Cassis tuberosa. No locality or color was given.

The two new species of the genus Vioa, V. dissociata and V. duvernoysii,
reported by Duchassaing de Fonbressin (1850) were described only in terms of
the gross morphology of their internal lobes and are thus unrecognizable. It
should also be noted that the name V. duvernoysii is preoccupied (V. duvernoysii
Michelin, 1847). The complicated taxonomic history of this name is discussed by
Vosmaer (1933).

In their monograph on the sponges of the West Indies, Duchassaing and
Michelotti (1864) included several new species of excavating sponges. They also
established a new genus, Euryphylle (written Euryphylla in the explanation of
the plates), for excavating sponges inhabiting corals. According to Topsent
(1888) this genus is a synonym of Cliona. The species descriptions by Duchas-
saing and Michelotti are too brief to be useful to subsequent workers in the
West Indian region. In addition, no reference is made to internal structures:
spicular morphology and dimensions or histology. Of the three new species,
Euryphylle dubbia and Vioa (misprinted Viva but subsequently corrected to
Vioa) strombi were considered by Vosmaer (1933) to be insufficiently described
and, thus, unrecognizable. V. strombi was found on Strombus gigas. The third
species, Euryphylle latens, was believed by Topsent (1889) to be, most probably,
his Cliona euryphylle.

Schmidt (1870) included two new species of his genus Papillina in a com-
munication on the sponges of the Atlantic. These species, both from Florida,
are Papillina arenosa and Papillina cribraria. De Laubenfels (1936a) stated that
P. arenosa is unrecognizable except that it belongs to the genus Cliona. P.
cribraria was placed in synonomy with Spheciospongia vesparia (Lamarck)
Marshall by de Laubenfels (1932) after he examined Schmidt’s type at the
Museum of Comparative Anatomy, Harvard University.

Vioa schmidti from Florida (Ridley, 1881) is not a clionid and should not be
confused with Cliona schmidti (Ridley) Topsent.

Topsent (1888) reported specimens of five species of clionids inhabiting shells
or corals from the Campeche Bank in the Gulf of Mexico. These sponges,
dredged during the cruise of the “Tabasco”, are: Cliona celata Grant, Cliona
vastifica Hancock, Cliona carpenteri Hancock, Cliona vermifera Hancock (from
the shell of the spiny oyster, Spondylus), and Cliona euryphylle (a new species
found in coral and in the shell of an undesignated species of Chama). In 1889

6 POSTILLA 161

Topsent enlarged his list of the clionids from the Campeche Bank to include
Cliona johnstoni (Schmidt) and Cliona subulata Sollas, in addition to the above
mentioned species. C. subulata was synonomized with Cliona viridis (Schmidt)
Gray by Topsent in 1891. It should be noted, however, that C. johnstoni was
later removed from the genus by Topsent (1900) and is not now considered to be
an excavating sponge. In the second part of his 1889 paper, Topsent reported C.
caribbaea Carter and Cliona labyrinthica Hancock inhabiting specimens of the
coral Porites from Point a Pitre, Guadeloupe, Leeward Islands, in the Lesser

Antilles.
Leidy (1889) believed that his species Cliona phallica might be a variety of

Cliona sulphurea Verrill and noted (1889, 1891) the similarity of C. sul/phurea to
European specimens of C. celata. C. sulphurea was put into synonymy with C.
celata by Topsent in 1891. Leidy’s C. phallica, from Florida, is a massive sponge
with tylostyles resembling those of C. su/phurea in their shape and size. It is in-
sufficiently described.

Verrill (1907) reported the presence of Cliona caribbaea Carter in Bermuda.
He also noted the existence of a massive form, with tylostyles but no micro-
scleres. Since he believed that this sponge might not be the gamma form of
Carter’s species, he proposed for it the provisional name Cliona sordida. {n the
same paper, Verrill also proposed a new genus, Heterocliona (type species,
Papillina cribraria Schmidt, 1870) for a massive or goblet-shaped sponge that
might have an excavating state when young. H. cribraria is perhaps also a
synonym of Spheciospongia vesparia.

De Laubenfels (1936a) stated that the genus Cliona is well represented in the
West Indian region and listed the following West Indian species: Cliona viridis,
Cliona phallica, Cliona subulata, and Cliona vermifera. This list is apparently
taken from the literature and not from collections made during the course of de
Laubenfels’ study. De Laubenfels described C. caribbaea in detail, and con-
sidered the possibility that it is a variety of C. celata. The validity of C. caribbaea
aS a Species is discussed in a subsequent section.

Only two species of clionids have been reported previously from Jamaica.
They are Cliona vermifera and Cliona viridis, both found in shallow water near
Port Royal on the south coast of the island (Hechtel, 1965). Hechtel also re-
ported the presence of Anthosigmella varians (Duch. & Mich.) de Laubenfels
in this area. From Bermuda we have another record of Cliona caribbaea and the
first description of Cliona lampa (de Laubenfels, 1950a). De Laubenfels (1949)
found Cliona vastifica and A. varians in the shallow waters of the western
Bahamas. He reported C. caribbaea from the Panama Canal Zone.

There have been two recent studies of the sponges of the Gulf of Mexico (de
Laubenfels, 1954a; Little, 1963). De Laubenfels reported the presence of Cliona
caribbaea and C. lampa. Little found Cliona truitti Old, C. lampa, C. celata, C.
vastifica, and C. viridis. However, the specimens reported as C. viridis were
misidentified and actually are Suberites undulatus George & Wilson (Hechtel,
1965).

Many of the reports of clionids from Florida are from faunal checklists. These
include C. vastifica from Apalachicola Bay, Florida (Pearse and Wharton, 1938);
C. celata from Boca Ciega Bay, Florida (Dragovitch and Kelly, 1964), St.

JAMAICAN EXCAVATING SPONGES Uf

George’s Sound, Florida (Menzel, 1956), and the west coast of Florida (Carter,
1885, as Raphyrus griffithsi); C. caribbaea from Florida Bay (Tabb and Mann-
ing, 1961).

Cliona celata has often been reported, as C. sulphurea, from the coast of
Louisiana in connection with the oyster industry there (Moore, 1899; Cary,
1906a,b, 1907). Hopkins (1956) identified four species of clionids (C. celata, C.
lobata, C. vastifica and C. truitti) from oyster shells of the Louisiana bays and
sought to determine the relative abundance of these species in terms of salinity.
His tentative identifications of species from Texas were based on external form
and the dimensions of the galleries. Hartman (1958) reported C. celata from
1.5 min Aransas Bay, Texas.

Previous Reports of Clionids and Related Sponges from the West Indies,
Florida, and the Gulf of Mexico
WEST INDIES
Jamaica:
Cliona vermifera Hancock, Hechtel, 1965, p. 61
Cliona viridis (Schmidt), Hechtel, 1965, p. 62
Bahamas:
Cliona vastifica Hancock, de Laubenfels, 1949, p. 20
Bermuda:
Cliona lampa de Laubenfels, 1950a, p. 110
Cliona caribbaea Carter, Verrill, 1907, p. 299; de Laubenfels, 1950, p. 108
Cliona sordida Verrill, 1907, p. 299
Guadeloupe:
Cliona caribbaea, Topsent, 1889, p. 49
Cliona labyrinthica Hancock, Topsent, 1889, p. 49
St. Vincent:
Cliona caribbaea Carter, 1882, p. 346
Undesignated locality within the West Indies:
Cliona millepunctata Hancock, 1849, p. 341
Euryphylle dubbia Duch. & Mich., 1864, p. 113
Euryphylle latens Duch. & Mich., 1864, p. 113
Vioa strombi Duch. & Mich., 1864, p. 113
Vioa duvernoysii Duchassaing, 1850, p. 27
Vioa dissociata Duchassaing, 1850, p. 27
FLORIDA
Atlantic coast:
Papillina arenosa Schmidt, 1870, p. 48
Papillina cribraria Schmidt, 1870, p. 48
Dry Tortugas:
Cliona caribbaea, de Laubenfels, 1936a, p. 155
Gulf coast:
Cliona vastifica, Pearse and Wharton, 1938, p. 365
Cliona celata Grant, Dragovich and Kelly, 1964, p. 78, Menzel; 1956, p. 1;
Carter, 1885, p. 207 (as Raphyrus griffithsi)
Cliona caribbaea, Tabb and Manning, 1961, p. 542

8 POSTILLA 161

GULF OF MEXICO

Texas:
Cliona celata, Hartman, 1958, p. 17

Louisiana:
Cliona celata, Hopkins, 1956, p. 49; as Cliona sulphurea by Moore, 1899,

p. 93, and Cary, 1906a, p. 21; 1906b, p. 50; 1907, p. 28

Cliona vastifica, Hopkins, 1956, p. 49
Cliona lobata Hancock, Hopkins, 1956, p. 49
Cliona truitti Old, Hopkins, 1956, p. 49

Florida, Gulf Coast:
Cliona lampa, de Laubenfels, 1953a, p. 540; Little, 1963, p. 57
Cliona caribbaea, de Laubenfels, 1953a, p. 540
Cliona truitti, Little, 1963, p. 57
Cliona vastifica, Little, 1963, p. 57
Cliona celata, Little, 1963, p. 57

Campeche Bank:
Cliona celata, Topsent, 1888, p 82; Topsent, 1889, p. 34
Cliona vastifica, Topsent, 1888, p 82; Topsent, 1889, p. 35
Cliona carpenteri Hancock, Topsent, 1888, p. 82; Topsent, 1889, p. 35
Cliona vermifera, Topsent, 1888, p. 82; Topsent, 1889, p. 35
Cliona euryphylle Topsent, 1888, p. 82; Topsent, 1889, p. 35
Cliona subulata Sollas, Topsent, 1889, p. 34

III. Descriptions of Jamaican Species of Excavating Sponges

Order HADROMERIDA

Family CLIONIDAE
Cliona schmidti( Ridley) Topsent

Vioa schmidtii Ridley, 1881, p. 130.

Cliona schmidti, Topsent, 1888, p. 87.

non Vioa schmidtii Carter, 1882, p. 348.

Vioa johnstonii var. Schmidt, 1870, pp. 5, 88.

non Vioa johnstonii Schmidt, 1862, pp. 78, 83, 86.
Vioa johnstonii Carter, 1882, p. 354.

Color. In life, eee to reddish purple. Underwater the sponge appears bright
lue to the d . After fixation in 10% neutralized formalin and transfer to 70%
after drying, it is dark purple. In acid alcohol (70% EtOH +

‘he sponge imparts a slight purplish tinge to the solution but retains

JAMAICAN EXCAVATING SPONGES 9

Substrates. Montastrea annularis (Ellis & Solander), M. cavernosa (Linnaeus),
Mussa angulosa (Pallas), Porites furcata Lamarck, Madracis mirabilis (Duchas-
saing & Michelotti), M. decactis (Lyman), Agaricia agaricites (Linnaeus) forma
agaricites, A. agaricites forma purpurea, Eusmilia fastigiata (Pallas), dead coral
and coral rock.

Depths. Collected from 15-47 m. This species has not been observed to occur
above 15 m in the study area, but a specimen inhabiting Mycetophyllia la-
marckana Milne-Edwards & Haime was collected from a depth of 58 m on
the Discovery Bay reefs (J.C. Lang, unpublished observations).

Material. Sixteen specimens from the reefs of Discovery Bay and Rio Bueno,
Jamaica, were studied.

Spicules. (1) Abundant tylostyles straight to slightly curved with globate to
subterminal heads and gradually tapering shafts. The heads are 7.9-13.8 um
(= micron) in maximum diameter. The tylostyles (Fig. 1A) are moderately long
and rather slender. Tylostyle length: 256 + 4.9 wm, 183-308 pm. Tylostyle width:
5.4 + 0.3 wm, 3.8-9.5 wm. N=8 specimens. Fifty spicules of each category were
measured for each specimen. (2) Numerous spirasters of two kinds, both of
which are spiny. The short, stout spirasters (Fig. IC) are straight to slightly
curved with prominent spines that are thorny in appearance and irregularly
distributed along the length of the axis. Length: 44.8 + 1.9 wm, 26.5-68.9 um.
Width: 5.8 + 0.1 um, 4.1-7.9 wm. N=8 specimens. The long thin spirasters
(Fig. 1B) have 7 to 10 turns to their spire. Their spines are more delicate in
appearance and are spirally arranged. Length: 73.9 + 1.9 wm, 60.9-100.7 um.
Width 2.0 + 0.04 wm, 1.6-3.2 »m. N=8 specimens. The width presented for
both categories of spirasters is that of the axis itself without the spines. The ab-
solute ranges of the lengths of these two groups of microscleres overlap some-
what. The means, however, are significantly different. Measurements made on
individual specimens show that the longer of the short, stout spirasters are also
somewhat thinner and rather resemble the long, thin spirasters in appearance.
These may not represent two distinct categories of microscleres.

Surface. The sponge does not encrust its substrate. Ostial and oscular papillae
are numerous and separate. Fusion of papillae has not been observed to occur.
The perforations in the coral skeleton or coral rock, through which oscular and
ostial papillae protrude, form very regular circles of rather variable size. The
oscules are shaped like truncated cones and the ostial papillae are low mounds
with sievelike surfaces. Both the oscular and ostial papillae are, even when
observed in place on the reef and fully expanded, rather low and close to the
surface of the substrate. Diameter of ostial perforations: 1.02 + 0.12 mm, 0.12-
2.46 mm. N=5 specimens. Diameter of oscular perforations: 1.83 + 0.30 mm,
0.80-2.40 mm. N=4 specimens. Some of the papillae, particularly the ostial
papillae, are quite small (less than 0.5 mm in diameter). These are, most prob-
ably,newly formed.

10 POSTILLA 161

Fig. 1. Spicules of Cliona schmidti (Ridley). A. Tylostyle. B. Thin spirasters. C.
Stout spirasters. Scale | (A) = 25 wm; scale II (B,C) = 10 pm.

Excavations. The sponge forms spherical or ellipsoidal galleries of rather irregu-
lar Outline. These galleries lie close to one another with only narrow remnants
of the coral skeleton between them. Diameter of the excavations: 1.17 + 0.06
mm, 0.49-2.64 mm (short diameter), 2.56 + 0.05 mm, 1.48-3.08 mm (long
ameter). N=8 specimens. The tissue of the sponge within the galleries is diapha-
10uU pon examination of freshly killed specimens, the tissue is seen to be
ppiied to the skeleton of the coral. The channels or tunnels leading

ithin the substrate to the surface openings are narrow and

JAMAICAN EXCAVATING SPONGES 11

Internal Structure. The tissue of the sponge is thin and loosely organized. Below
the exopinacoderm, the mesohy! has rather large elliptical lacunae (190 4m by
90 um). The eurypylous choanocyte chambers are 20-25 um in diameter. Num-
erous small (10-14 xm) pigment cells containing granules that stain darkly with
hematoxylin are scattered throughout the tissue. These cells appear capable of
assuming a variety of shapes (see Topsent, 1900) and, due to their intrinsic
violet coloration, are also clearly visible in unstained sections. Tylostyles and
short, stout spirasters lie together in profusion, particularly in the basal region
of the papillae. Long, thin spirasters are confined to the choanosome and are
most frequently found around the periphery of canals. The short, stout spiras-
ters are the most abundant skeletal elements. Throughout the tissues, groups of
highly eosinophilic, globule-containing cells (12-15 ~m in diameter) with a
prominent nucleolus are both numerous and conspicuous. These cells, not de-
scribed by any previous investigators dealing with C. schmidti, are, perhaps,
algal.

Taxonomic Discussion. Specimens of C. schmidti were first reported by Schmidt
(1870) as a variety of his Vioa johnstonii. Carter (1879) suggested that the two
are separate species having in common only their striking purple coloration.
Ridley (1881) designated Schmidt’s (1870) variety as V. schmidtii. V. johnstonii
is not an excavating sponge and was transferred in 1898 to the genus Coppatias
(=Jaspis) by Topsent. In 1882 Carter unfortunately decided upon V. johnstonii
as the name for the 1870 variety and used V. schmidtii for Schmidt’s original
species. Ridley (1884) included Carter’s 1882 reference in his synonomy but did
not mention the discrepancy. Topsent transferred Ridley’s V. schmidtii to the
genus Cliona in 1888, but Lendenfeld (1897) continued to refer to this species
as Vioa schmidtii. A carmine-colored excavating sponge from Australia was
designated Vioa johnstonii by Carter (1886). This sponge was described by
Carter as possessing tylostyles and spirasters. Tylostyle measurements were not
given; the tylostyles were neither described nor figured. The spirasters, for which
measurements were given, are quite short (42 4m). Topsent (1900) doubted that
this is a specimen of C. schmidti (Ridley). The presence of C. schmidti in
Australia is therefore questionable.

Geographical Distribution. Mediterranean: Monaco, Topsent, 1934, p. 13;
Bonifacio (Corsica), Topsent, 1932, p. 574; Gulf of Gabes (Tunisia), Topsent,
1894, p. 43; ibid., 1934, p. 73; Gulf of Policastro (Italy), Sara, 1963, p. 209;
Banyuls (France), Topsent, 1892, p. xvii. Adriatic: Rovigno, Volz, 1939, p. 16;
Lesina, Lendenfeld, 1897, p. 74; Cattaro, Schmidt, 1870, p. 5; Tremiti Island,
Sara, 1961, p. 38. Red Sea: Topsent, 1906, p. 570ff. Indo-Pacific: Eagle Island
(Amirante Islands), Ridley, 1884, p. 589; ?Port Western (Australia), Carter,
1886, p. 458; ?Komebail Lagoon (Palaus), de Laubenfels, 1954, p. 217.

Previous Descriptions and Discussion. The type of C. schmidti was described
from the Adriatic and this species has been reported frequently from the shallow
waters of both the Adriatic and Mediterranean seas. Although the Jamaican
specimens described above were all collected from relatively deep water (15-

12 POSTIELA I6]

47 m), they agree quite well with those from the Adriatic and Mediterranean.
Volz (1939), for example, gave the following dimensions for the spicules of his
specimens of C. schmidti: mean length and width of tylostyles, 210-280 4m by
4 wm; mean length and width of the long, thin spirasters, 70 wm by 2.4 wm.
Topsent’s (1900) description of the spicular morphology and dimensions, sur-
face morphology, papillary dimensions, and histology of this species also agrees
quite well with that presented above for the Jamaican specimens. Previous in-
vestigators have all been struck by what Schmidt called the “magnificent violet”
color so characteristic of this species. De Laubenfels (1954) reported the presence
of C. schmidti from the shallow waters of the west central Pacific. His speci-
mens, however, were brown in life with tylostyles 260 zm by 3 »m in length and
spirasters 32-42 xm in length. Some of the shorter spirasters have a spiraled
shaft. De Laubenfels stated that they do not appear to be characteristic of the
species C. schmidti. Most probably de Laubenfels’ specimens are not C.
schmidti.

Cliona vermifera Hancock

Cliona vermifera Hancock, 1867, p. 239.
Vioa vermifera, Lendenfeld, 1897, p. 80.

Color. In life, red to red orange. Shallow water specimens (1 m) will appear
red orange to the diver while specimens from deeper water (18 m and more) will
appear brown. After fixation in 10% neutralized formalin and transfer to 70%
ethyl alcohol the sponge is dark orange yellow. Dried specimens are pale yellow.

Substrates. Agaricia undata (Ellis & Solander), Helioseris cucullata (Ellis &
Solander), dead coral (of the plate-like Agaricia- Helioseris type), and coral rock.

Depths. Collected from I-2 m and 18-52 m. The sponge has not yet been
collected from intermediate depths in the study area.

Material. Four specimens from the reefs of Discovery Bay, Jamaica, were
studied.

Spicules. (1) Abundant tylostyles (Fig. 2A, B) exhibiting a rather wide range of
lengths and widths (141-387 wm by 5.0-10.2 wm) for four specimens. The
shorter tylostyles have, for the most part, stout, curved, gradually tapering
shalts, and subterminal heads with a maximum diameter of 9.0-11.2 pm.
lylostyle length: 175 + 3.1 wm, 152-187 wm. Tylostyle width: 8.9 + 0.22 um,
5.1-9.9 um. N=4 specimens. The longer tylostyles are mostly thinner and
straight, tapering toward the pointed end. The head, with a maximum diameter

.2-10.8 um, may be round, slightly elongate or subterminal. Tylostyle
length: 287 + 2.6 wm, 195-352 um. Tylostyle width: 6.1 + 0.1 wm, 5.5-7.7 pm.

* specimens. The two categories of tylostyles described above are not com-

JAMAICAN EXCAVATING SPONGES 13

Fig. 2. Spicules of Cliona vermifera Hancock. A. Tylostyle. B. Tylostyles, head end
enlarged. C. Spirasters. Scale 1 (A) = 25 ym; scale Il (B,C) = 5 um.

pletely discontinuous; there are intermediate forms. (2) Numerous smooth,
“worm-like” (Carter, 1867, p. 239) spirasters (Fig. 2C) with blunt, rounded ends.
Some possess a slight swelling near the central region, but the majority are of a
uniform width. Almost all of the spirasters are multiply bent and have from
2-5 turns. A few are arched rather than undulating. Spiraster length: 43.8 +
1.2 pm, 31.8-52.7 wm. Spiraster width: 4.0 + 0.2 wm, 2.2-4.7 wm. N=4 speci-
mens. Each of the four specimens examined possessed some extremely thin
spirasters (less than 2 xm in diameter) of normal length. These are presumed to
be young stages. Fifty spicules in each category were measured for each
specimen.

14 POSTILLA 161

Surface. The numerous scattered ostial and oscular papillae (Fig. 20) are
separate. They form no distinct pattern on the surface of the substrate. Occa-
sional fusion of two ostial papillae may be observed, but the sponge does not
tend to overgrow its substrate. Ostial papillae are sievelike in appearance, bear-
ing numerous small ostia. The oscules are shaped like truncated cones. The
perforations through which the oscular and ostial papillae protrude are circles,
regular and smooth in outline, and are within the same size range. Diameter of
ostial perforations: 1.04 + 0.04 mm, 0.68-1.48 mm (absolute range).
specimens. Diameter of oscular perforations: 1.08 + 0.04 mm, 0.74-1.54 mm.
N=3 specimens.

Excavations. Relatively large galleries not quite completely filled with sponge
tissue in the form of rounded to elongated lobes interconnected by numerous
slender cylindrical stems. The excavations (Fig. 19) are discrete rather than con-
fluent. This sponge has been found most frequently during the course of the
present study in flattened, platelike species of agariciids (Agaricia undata,
Helioseris cucullata) having a thickness of approximately 7 mm or less. In such
substrates, the excavations extend nearly from the upper to the lower surface of
the coral, leaving very little of the skeleton remaining at either surface. Adjacent
galleries are separated by thin remnants of coral and tend to be lined up, one
after another, forming a row. Diameter of spherical excavations: 2.64 + 0.07
mm, |.11—4.30 mm (absolute range). N=4 specimens. Diameter of the ellipsoidal
excavations: 1.64 + 0.11 mm, 0.98-2.21 mm (short axis); 3.48 + 0.09 mm,
2.54-5.56 mm (long axis). N=4 specimens. Width of the substrate remaining be-
tween adjacent galleries: 0.97 + 0.10 mm, 0.82-1.20 mm. N=4 specimens.
Diameter of interconnecting stems: 0.32 + 0.08 mm, 0.26-0.51 mm. N=4 speci-
mens. The channels leading from the galleries within the substrate to the sur-
face openings are quite short, about 1-2 mm in length. The degree of excavation
of the substrate is usually quite extensive. When an infected coral is viewed in
longitudinal section, a row of exposed galleries may be seen extending along the
entire length of the cut surface.

Internal Structure. The relatively dense tissue contains numerous tylostyles. The
smooth spirasters are found mainly in the basal region of papillae. Numerous
eosinophilic, granular cells, 9-12 zm in diameter, are scattered throughout the
internal lobes. Choanocyte chambers are 18 to 20 xm in diameter.

Taxonomic Discussion. Many clionid species descriptions, particularly those of
the earlier workers who often had only dried specimens to examine, are based
largely on spicular morphology and dimensions. C. vermifera appears to be the
'y described clionid having a spicular complement composed solely of tylo-
nooth, stout spirasters. Consequently, this species has had an un-
nplicated taxonomic history. It was reported by Lendenfeld (1897)
latic Sea under the name Vioa vermifera. All previous and sub-

species have been as C. vermifera.

JAMAICAN EXCAVATING SPONGES INS)

Geographical Distribution. Tropical Atlantic: Gulf of Campeche (Mexico),
Topsent, 1889, p. 35; Port Royal (Jamaica), Hechtel, 1965, p. 60. Mediterranean:
Gulf of Naples, Vosmaer, 1933, p. 565. Adriatic: Lesina, Lendenfeld, 1897,
p. 80; Rovigno, Volz, 1939, p. 18. Indo-Pacific: Tuamotu and Gambier archi-
pelagos, Topsent, 1933, p. 565.

It should be noted that Volz (1939) was mistaken in stating that C. vermifera
was found among a collection of sponges from the West Indies and Acapulco
described by Carter (1882). C. vermifera appeared in Carter’s 1882 communica-
tion only as one of a group of sponges he considered as belonging to the family
Suberitidae. His purpose in compiling a list of these sponges was to “show that
the pin-like skeletal spicule is often accompanied by a spinispirular or other
flesh spicule, as well as often without it” (Carter, 1882, p. 354.)

Discussion. C. vermifera has been rather infrequently reported. Topsent
(1896) found isolated smooth, stout spirasters reminiscent of those of C. vermi-
fera and was thus tempted to include this species in his list of Hadromerina
from France. In his comprehensive study of 1900, however, he concluded that
these spirasters were most likely from Spiroxya heteroclita Topsent. Lendenfeld
(1897) and Vosmaer (1933) each reported only one specimen, but Volz (1939)
found the species to be quite common in the shallow waters in the vicinity of
Rovigno. The present study indicates that the species is not common at
Discovery Bay, on the north coast of Jamaica. Hechtel (1965) reported only one
specimen of this species, from shallow water, at Port Royal on the south coast.
Unfortunately, the type locality of C. vermifera is not known. Hancock (1897)
stated that he examined two specimens inhabiting a species of Chama, a eula-
mellibranch having a very wide geographical distribution. It is also unfortunate
that Hancock did not report the range of tylostyle lengths for his type. He gave
the tylostyle length as 254 wm. Subsequent workers have found a rather wide
range of tylostyle lengths, widths and shapes in individual specimens. Volz
(1939), however, reported as megascleres only rather small tylostyles (mean of
175 wm) with subterminal heads; he did not give the range of lengths. It should
also be noted that the spirasters of specimens from the Mediterranean and
Adriatic (Vosmaer, 1933 and Volz, 1939 respectively) are not as pronounced ly
undulating as those described by Hancock and other investigators. However,
since the other characters, as described by Topsent, Lendenfeld, Vosmaer,
Volz, and Hechtel are all quite similar to those of Hancock’s C. vermifera, there
seems to be no reason to doubt that we are dealing with one widely distributed
species. The West Indian specimen figured by Topsent (1900, Plate I, fig. 4)
also possesses the thin spirasters found during the present study. Topsent (1889,
1900) did not, however, mention the presence of a swelling near the central re-
gion of the spirasters. This swelling does occur on a few of the spirasters of
Hechtel’s (1965) specimen. The granular, spherical cells described by Hechtel
(1965, p. 60) were also found in the specimens from Discovery Bay.

16 POSTILLA 161

Cliona janitrix Topsent
Cliona janitrix Topsent, 1932, p. 575.

Color. In life, dull yellow. Underwater, the sponge will appear to be almost
colorless to the diver. After fixation in 10% neutralized formalin and transfer
to 70% ethyl alcohol it is very pale yellow, almost colorless. Dried specimens
are pale yellow.

Substrates. Acropora cervicornis (Lamarck), Madracis mirabilis (Duch. &
Mich.), Porites furcata Lamarck, P. porites (Pallas), and dead coral (cylindrical,
branched pieces).

Depths. Collected from 15-28 m.

Material. Five specimens from the reefs of Discovery Bay and Rio Bueno,
Jamaica, were studied.

Spicules. (1) Abundant, slightly curved, moderately stout tylostyles (Fig. 3).
The heads are regular in shape having either subterminal or terminally rounded
knobs. Maximum knob diameter is 10.9-12.0 wm. The shaft tapers gradually
toward the pointed end. This end of the spicule, however, is most irregular in
its form. Beginning approximately 30 um before the termination of the shaft of
the majority of the tylostyles there is a series of 2-3 notches, each one diminish-
ing the diameter of the shaft. The pointed tips of the spicules may be irregularly
crenulate (Fig. 3). The remainder of the tylostyles taper evenly to about 10 pm
above their termination. They then end abruptly in a point of greatly dimin-
ished diameter. An enlargement of the axial canal may often be observed in
the head region. Tylostyle length: 195 + 6.1 um, 134-221 pm. Tylostyle width:
9.2+0.4 wm, 7.8-11.4 wm. N=5 specimens. Occasional oxeiform spicules occur
with both ends showing one or more abrupt reductions in diameter.

Surface. Very small, scattered ostial and oscular papillae not forming any dis-
tinct pattern on the surface of the substrate. Papillary fusion has not been
observed to occur; the sponge does not overgrow the substrate. Both the ostial
and oscular papillae are low, close to the surface and not at all prominent. Fully
expanded oscules present the appearance of short cylinders. Ostial papillae are
lacy, bearing numerous small ostia. The perforations through which the ostial
and oscular papillae protrude are quite circular in their outline. Diameter of
the ostial perforations: 0.58 + 0.02 mm, 0.37-0.98 mm. N=4 specimens. Diame-
ter of the oscular perforations: 0.70 + 0.03 mm, 0.43-0.98 mm. N=4 specimens.

Excavations. The large, spherical to roughly cylindrical galleries are completely
tilled with the soft, rather delicate tissue of the sponge. Adjacent lobes are con-
nected by lattened stems of varying lengths. Interconnecting stems have not
been observed between all the lobes in the same substrate. During the course
of the present study, the sponge has been found to inhabit only branched corals

JAMAICAN EXCAVATING SPONGES Le,

Fig. 3. Tylostyles of Cliona janitrix Topsent. Scale = 20 wm.

of the genera Porites, Madracis and Acropora. \n such substrates it may exca-
vate, and completely remove, the central axis of the coral branch. This 1s
particularly true of the specimens inhabiting Madracis mirabilis, since branches
of this coral are of rather small diameter (usually less than 5 mm). When a coral
infected in such a manner is viewed in cross section, a mass of dull yellow tissue
may be seen occupying the center of the branch. The channels leading from the

18 POSTIEEATG6I

galleries to the surface papillae are quite narrow (approximately 0.5 mm in
diameter) and relatively long, averaging 3-4 mm in length. Diameter of the
excavations: 5.1 + 0.2 mm, 3.5-6.0 mm. N=5 specimens.

Geographical Distribution. Mediterranean: Bonifacio (Corsica), Topsent, 1932,
Dp: SS:

Previous Descriptions and Discussion. This species has been reported only
once previously (Topsent, 1932). It is listed in Volz’s (1939) key as one of the
Mediterranean clionids not found during the course of his comprehensive study
of the boring sponges at Rovigno. Subsequent workers in the Mediterranean
and Adriatic have not reported the presence of C. janitrix in these seas (Russ
and Riitzler, 1959: Riitzler, 1965, 1966; Sara 1959, 1961, 1963; Vacelet, 1960),
nor was the sponge reported by Vacelet (1961) in his study of the Demospongiae
of Bonifacio, the type locality.

Topsent’s specimens are from large fragments of a rather thick oyster valve.
Neither the species of oyster nor the depth from which it was collected were
presented. The spiculation of his specimens is composed solely of tylostyles.
These tylostyles are very close to those of the Jamaican specimens in both their
form (fig. vii, p. 575) and their dimensions (180-210 um by 8-12 wm). Topsent
stated that both surfaces of the substrate were covered with foreign objects
rendering observation of the papillae impossible. The papillae were described
merely as small and of ordinary construction; the dimensions were not pre-
sented. The galleries (2-3 mm in diameter) excavated by Topsent’s species are
rounded, spacious and not compactly crowded together.

Cliona lampa de Laubenfels
Cliona lampa de Laubenfels, 1950a, p. 110.

Color. In life, vermilion. Underwater, the sponge appears brown orange to
the diver. After fixation in 10% neutralized formalin and transfer to 70% ethyl
alcohol it is colorless. Dried specimens are pale brown.

Substrates. Acropora cervicornis (Lamarck), A. palmata (Lamarck), Agaricia
agaricites (Linnaeus) forma agaricites, A. agaricites forma purpurea, Montas-
trea annularis (Ellis & Solander), Solenastrea bournonii Milne Edwards &
Haime, Porites astreoides Lesueur, encrusting coral rock and dead coral.

Depths. Collected from 0.5-36 m.

Material. Twenty-eight specimens from the reefs of Discovery Bay, Jamaica,

latively few tylostyles (Fig. 4A) with long, thin, straight shaft

a point. The terminal knobs are round to oval with a

JAMAICAN EXCAVATING SPONGES 19

4
e
3
Ba
%
«te
<a
i
4
z
4
ia
4
x
a
—q
;

Fig. 4. Spicules of Cliona lampa de Laubenfels. A. Tylostyle. B. Microspined oxea.
C. Spiny microrhabd. Scale I (A) = 25 pm; scale Il (B,C) = 5 wm.

maximum diameter of 4.9-8.2 um. Tylostyle length: 280 + 4.1 um, 236-388 pum.
Tylostyle width: 5.1 + 0.1 mm, 4.0-7.1 um. N=7 specimens. (2) Abundant
microspined oxeas (Fig. 4B) often exhibiting a central swelling or possessing
more prominent spines in the central region. Oxeas taper gradually from the
mid region to the pointed ends. Some oxeas are straight. Oxea length: 79.1 +
2.1 am, 60-116.3 wm. Oxea width: 3.1 + 0.1 pm, 2.1-4.3 wm. N=7 specimens.
It should be noted that oxeas from shallow water specimens (0.5-3 m) were
slightly larger (90.5 4m in length, mean of 3 specimens) than those from deeper
water specimens (15 m or more). The dimensions of the other two categories of
spicules remained constant regardless of depth. (3) Abundant spiny microrhabds
(Fig. 4C) that are always straight. Although they are quite small and thin, their
spines are more prominent than those of the oxeas. The microrhabds are the
most abundant skeletal element. Microrhabd length: 15.2 + 0.3 pm, 9.2-22.1
um. Microrhabd width: 2.0 + 0.1 ym, 1.1-2.6 wm. N=7 specimens. Both the

20 POSTILLA I61

oxeas and microrhabds are spined throughout their entire length. Fifty spicules
of each category were measured for each specimen considered.

Surface. Ostial and oscular papillae are separate and abundant. Although fu-
sion of two ostial papillae may occasionally be observed, the sponge does not
tend to encrust its substrate (see Discussion). The perforations in the substrate
through which oscular and ostial papillae protrude are roughly circular in out-
line. The diameter of the perforations is quite variable. When fully expanded,
the oscules (Fig. 17) appear as short cylinders projecting from the surface of the
substrate. The ostial papillae are also slightly raised up above the substrate and
present a very lacy appearance. Diameter of the ostial perforations: 0.72 + 0.06
mm, 0.18-1.48 mm. N=16 specimens. Diameter of the oscular perforations: 1.20
+ 0.14 mm, 0.25-3.44 mm. N=15 specimens. The diameter of the papillary
perforations appears to be correlated with the age of the sponge and, thus, also
with the extent of the excavations. When the sponge is maintained in the labo-
ratory, the oscular papillae are frequently observed to be surrounded by tiny
chips of calcium carbonate. The expulsion of these chips from the excurrent
openings may occasionally be seen.

Excavations. The spherical to ellipsoidal galleries (Fig. 22), separated from each
other by narrow remnants of the coral skeleton, form a honeycombed pattern
when viewed in cross section. These galleries (Fig. 18), each completely filled
with a mass of soft sponge tissue, are occasionally confluent. The internal lobes
of the sponge are interconnected by slender stems (approximately 0.1-0.2 mm
in diameter). Diameter of the spherical excavations: 1.05 + 0.05 mm, 0.31-1.72
mm. N=5 specimens. Diameter of the ellipsoidal excavations: 1.08 + 0.10 mm,
0.43-2.03 mm (short axis); 1.58 + 0.10 mm, 0.74-3.08 (long axis). N=8 speci-
mens. Relatively short, wide channels leading to the surface papillae originate at
irregular intervals, from individual chambers within the network of galleries.
Diameter of the channels: 1.21 + 0.05 mm, 1.02—1.48 mm. Length of the chan-
nels: 3.42 + 0.09 mm, 3.12—-4.23 mm. N=8 specimens. In Acropora cervicornis,
the excavations may occupy the entire central axis of a branch, to a length of
12-15 cm. In the more massive species of corals the excavations of this sponge
are confined to the more superficial layers. This may be related to the channel
length of the species.

Internal Structures. The loosely organized tissue of the sponge, apparently lack-
ing species-specific spherular cells, has oxeas strewn in confusion. Although
some spiny microrhabds are present in the internal lobes, they are localized in
dense bands within the papillae. The relatively scarce tylostyles also contribute
to the framework of the papillae. Choanocyte chamber diameter is 14.5-17.4
um.

Geographical Distribution. Tropical Atlantic: Bermuda, de Laubenfels, 1950a,
p. 110; Gulf of Mexico, de Laubenfels, 1953a, p. 541; Tampa (Florida), Little,

1963, p. 57.

Cliona lampa is apparently confined in its distribution to tropical Atlantic
America including the West Indies, Florida and the Gulf of Mexico.

JAMAICAN EXCAVATING SPONGES 21

Previous Descriptions And Discussion. Although I have placed the specimens
described above in the species C. /ampa, there do exist some noteworthy dif-
ferences between these specimens and de Laubenfels’ species. C. lampa, as
originally described by de Laubenfels (1950a) from the shallow waters of Ber-
muda, “covers the entire surface of the mineral in which it lives” and is not
confined to galleries but “penetrates interstices in all directions” (p. 111). De
Laubenfels did, however, report the existence of “strands of continuous sponge
protoplasm buried in the rocks” that “may correspond to ordinary Cliona gal-
lenies/(1950a; jp: 111):

In the subsequent descriptions of C. /Jampa from the Gulf of Mexico (de
Laubenfels, 1953a), Tampa, Florida (Little, 1963), and Bermuda (Neumann,
1966), reference was also made to the ability of this species to permeate its sub-
strate. In addition to this, Neumann (1966) described labyrinthine galleries one
or more millimeters in diameter for specimens from Bermuda. Thus far, this
habitus of excavating galleries and permeating the substrate has been described
for specimens found in aeolian limestone (de Laubenfels, 1950a), Siderastrea
siderea (Little, 1963), dead coral, porous calcarenite and dense, well-cemented
limestone (Neumann, 1966).

The Jamaican specimens discussed above exhibit no tendency to encrust or
to permeate the substrate. The sponge is always confined solely to the honey-
comb excavations, the channels leading to the surface, and the papillae.

From de Laubenfels’ original description, the spicule complement consists
of straight tylostyles measuring 3 by 150 »m to 3 by 210 wm; microspined oxeas,
2 by 70 pm, and microspined streptasters (referred to above as spiny micro-
rhabds), | by 10 pm. The measurements given by Little are somewhat larger.
Tylostyles: 153-2/0-240 um by 2-3.9-6 wm. Oxeas: 77—-92.0-105 ym by 2-2.4-3
pm. Streptasters: 5—8./—-13 wm by 1-/.9-2.2 wm. Although the microscleres des-
ignated “streptasters” by de Laubenfels are straight and rodlike, the term strep-
taster was originally employed by Sollas (1888) for a category of spiral micro-
scleres to be distinguished from euasters with spines radiating from a central
point. This term has since been used in various other ways (see Reid, 1970),
notably by de Laubenfels (1950a, 1955) to refer to a “straight rod with long
spines or rays” (1955, p. E30). Following the terminology of Sollas, microrhabd
has been substituted here for de Laubenfels’ “streptaster”. The tylostyles of the
Jamaican specimens are larger and more robust, and spination of the strep-
tasters is more pronounced than those in de Laubenfels’ original description.
Although the spicules are, in general, larger than those of Little’s specimens,
the oxeas are somewhat smaller. De Laubenfels did not mention the presence
of a central swelling on the oxeas. However, examination of the spicules from
a portion of his holotype (BMNH register number 1948.8.6.45) has revealed
that many, but not all, of the oxeas do exhibit a central swelling. The same
condition has been noted for the Jamaican specimens.

Neither de Laubenfels (1950a, 1953a, b) nor Little (1963) gave descriptions
or dimensions of the pores or oscules. Neumann (1966) stated that the oscules
are often 3-4 mm in diameter. This figure is somewhat larger than the mean
oscular diameter found for the Jamaican specimens. However, a few of these
specimens had oscules measuring 2—3 mm in diameter.

oy) POSTILLA 161

Nothing has been reported previously concerning the internal structure of
this species.

De Laubenfels gave the color of C. /Jampa in life as “brilliant red, tending
toward the vermilion” (1950a, p. 110); Little, “a distinctive brick red” (1963, p.
57); and Neumann, “brilliant vermilion” (1966, p. 102). The Jamaican specimens
are vermilion. De Laubenfels further stated that after preservation in alcohol
C. lampa_ is “nearly white” and that after drying specimens are “slightly
brownish but very pale” (p. 110). Similar conditions are found in the Jamaican
specimens, as has been noted above.

According to de Laubenfels (1950a), C. Jampa is most abundant in areas with
strong currents. “For example, at the inlet to Harrington Sound where the water
is eternally rushing, either in or out, there are huge areas of this sponge, areas
of hundreds of square meters” (p. 110). The Jamaican specimens exhibit no
particular preference for areas with fast water movement.

In summary, the spicular morphology of de Laubenfels’ C. /ampa and that
of the Jamaican specimens examined during the course of the present study is
certainly extremely similar. The spicular dimensions of the latter (except for
the oxeas) are somewhat larger. Although the Jamaican specimens do not per-
meate or encrust their substrates, the boring habit is also similar. Further col-
lections from Bermuda are needed to establish the depth range of C. lampa
there. It would also be of great interest to determine whether there exist speci-
mens of this species from Bermuda that do not encrust or permeate their
substrates. At this time, however, it does not appear that there is sufficient
justification for considering the Jamaican specimens as belonging to a new
species.

Cliona caribbaea Carter

Cliona caribbaea Carter, 1882, p. 346.

(non) Cliona caribbaea, de Laubenfels, 1936a,b; 1950a; 1953a,b; Wells, Wells
and Gray, 1960, p. 228.

Cliona viridis, Topsent, 1900, p. 84 (partim).

Cliona viridis, Hechtel, 1965, p. 61.

Cliona viridis var. caribbaea, Topsent, 1932, p. 563.

Color. In life, the papillae are yellow; the internal lobes are dull yellow. Under-
water, the sponge appears dull yellow to the diver. After fixation in 10% neu-
tralized formalin and transfer to 70% ethyl alcohol, it is dark green.

Substrate. Porites porites (Pallas).
Depths. Collected from 1-2 m.

Viaterial. Four specimens were collected from the lagoon zone (Kinzie, 1970)
iscovery Bay reefs. Only small pieces of these specimens were preserved
lent spicule preparations.

JAMAICAN EXCAVATING SPONGES 23

Fig. 5. Spicules of Cliona caribhbaea Carter. Specimen from Jamaica. A. Tylostyles.
B. Spiraster. Scale 1 (A) = 30 wm; scale II (B) = 5 wm.

Spicules. (1) Abundant, very long, rather stout tylostyles (Fig. 5A). The shaft,
usually slightly curved, but sometimes straight, has its greatest diameter at the
midpoint and begins to taper at a point approximately two-thirds of the dis-
tance from the head to the tip. The head is elongated, elliptical or, occasionally,
spherical. The variation in the form of the knob is not great; it is never sub-
terminal. Maximum diameter of the knob is 12.0-16.1 wm; length is 16.0-18.4
pm. Immediately behind the head of a few tylostyles there may be a secondary
annular swelling. Occasionally this swelling occurs further down the shaft.

14 POSTILLA 161

Tylostyle length: 394 + 2.4 wm, 361-451 pm. Tylostyle width: 10.2 + 0.2 «um,
9.4-12.1 um. N=3 specimens. (2) Abundant spiny spirasters (Fig. 5B) usually
having 2-4 turns. The spines, thornlike in appearance, are spiralled around the
axis of the spiraster. Some are very prominent, 3 wm in height. Both ends of the
axis bear prominent spines that bifurcate or trifurcate. There are a few rather
thin (approximately | ~m in diameter), almost straight, spirasters with small
spines. Spiraster length: 38.3 + 1.1 wm, 27.3-41.8 pm. Spiraster width:
1.5+0.04 pm,!.0-2.2 »m. N=3 specimens.

Surface. The relatively large, scattered ostial and oscular papillae, lying close
to the surface of the substrate, are numerous and separate. No papillary fusion
was observed. The sponge exhibits no tendency to overgrow the substrate and
enter the massive or gamma stage. Measurements of papillary perforations
were not made.

Excavations. The galleries are large, cylindrical, ragged in their outline, and
not completely filled with the rather tough tissue of the sponge. In the study
area, the sponge has thus far only been observed to inhabit branches of the
coral P. porites (dead basal region of living colonies). In this substrate, C. carib-
baea produces excavations that completely remove the central axis of the in-
fected branch. The internal lobes of the sponge often bifurcate. Diameter of
the excavations: 4.2 mm (mean of three specimens), 3.7-5.8 mm (absolute range
of three specimens). Short, wide channels connect the internal lobes with the
surface papillae.

Geographical Distribution. Tropical Atlantic: Point a Pitre (Guadeloupe), Top-
sent, 1889, p. 49; St. Vincent, Carter, 1882, p. 346; Bermuda, Verrill, 1907, p.
299; Port Royal (Jamaica), Hechtel, 1965, p. 61 (as Cliona viridis).

This species (sensu Carter, 1882) has been reported only from the West Indian
region.

Discussion. The question of the validity of Carter’s species, Cliona caribbaea,
presents us with a complex situation. The list of “Previous Reports” above
shows that since its original description in 1882 there have been eight reports of
C. caribbaea from the West Indies, Florida and the Gulf of Mexico. Topsent
(1891) placed C. caribbaea and C. subulata (also reported from the West Indies)
in synonymy with C. viridis and later (1900, Pl. VII, fig. 2b) figured spicules of
his Leeward Islands specimens of C. caribbaea as an example of the spiculation
of C. viridis. Hechtel (1965) also considered C. caribbaea and C. subulata to be
synonyms of C. viridis which he reported from Jamaica.

From the original description of C. swbulata (Sollas, 1878, p. 65), we know
only that this species, along with C. mucronata Sollas and C. ensifera Sollas,
was found in excavations in a “patch of Melobesia” that was “associated with
Isis” tp. 65), a gorgonian from an undesignated locality. The spicules, rather
thin tylostyles with globose knobs, and spiny spirasters were figured (P1. II, figs.
26, 27) but not described; no dimensions were given. Because of the brief nature

JAMAICAN EXCAVATING SPONGES pe)

of the original description, there does not appear to be sufficient evidence to
consider C. caribbaea to be a junior synonym of C. subulata.

C. viridis, originally described from the Adriatic by Schmidt, is quite common
in the relatively shallow waters of the Adriatic and Mediterranean Seas (T op-
sent, 1900; Volz, 1939; Sara, 1961, 1963, 1965a). The type specimen of C.
viridis, rather briefly described by Schmidt in 1862, possessed tylostyles with
subterminal knobs that are quite distinctive. Tylostyles with the same spatulate
heads were observed in a Mediterranean specimen (YPM No. 2896) examined
by the present investigator. The spicules of this specimen, both the tylostyles
and spirasters, were also quite distinctive. The morphology and dimensions of
these spicules are given in Table |. Topsent’s (1900) specimens of C. viridis ex-
hibit a great variety in the shape of the head region (it may be subterminal,
elongated, elliptical or globose), the color of the sponges in life, the form and
length of the spirasters, the arrangement and length of the spiraster spines and
the size of the papillae. He apparently included his West Indian specimens of
C. subulata and C. caribbaea in his description of C. viridis. Topsent (1900) also
described three types of spherular cells that are species-specific for C. viridis.
He stated that these cells are also present in West Indian specimens of C. sub-
ulata and C. caribbaea. Histological studies of the Jamaican sponge are needed.

The Mediterranean specimens of C. viridis exhibit a very pronounced ten-
dency to overgrow the substrate and enter a thickly encrusting stage (Topsent,
1900, and Hartman, pers. comm.). Aside from the gamma stage specimens re-
ported by Verrill, de Laubenfels and Wells et al. (discussed below), C. carib-
baea shows no tendency toward papillary fusion and has not been observed to
encrust or overgrow the substrate (Carter, 1882; Topsent, 1889; Hechtel, 1965;
Discovery Bay specimens).

The three specimens collected during the course of the present study agree
quite well with those described by Hechtel (1965) under the name C. viridis.
Hechtel also noted the presence of a secondary annular swelling behind the head
region of the tylostyles.

Carter (1882) believed that the characteristics of his species were very close to
those of C. celata, but that C. caribbaea could be distinguished from C. celata
by the spherical head of its tylostyles (“skeletal spicules’) and the presence of
spirasters (“flesh spicules”). Dr. W.D. Hartman kindly provided a spicule slide
prepared from a fragment of Carter’s type deposited in the City of Liverpool
Museums. See Figure 6.

As mentioned previously, de Laubenfels (1936a,b, 1950a, 1953a,b) did not
report the presence of spirasters in his specimens. The length of the tylostyles
is also somewhat less than that of the type. De Laubenfels stated that the sponge
may often reach the massive or gamma stage. The specimen he described from
Dry Tortugas came from deep water (70 m) and inhabited extremely wide gal-
leries not completely filled by the tissue of the sponge. It is doubtful whether
any of these specimens (from Dry Tortugas, Panama, Bermuda and the Gulf of
Mexico) are actually attributable to the species described by Carter.

Verrill (1907) stated that young individuals of C. caribbaea from Bermuda
“excavate extensive and irregular cavities in shells and corals, especially Porites”

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JAMAICAN EXCAVATING SPONGES 27

Fig. 6. Spicules of Cliona caribbaea Carter. Holotype from St. Vincent, W.1. A.
Tylostyles. B. Spiraster. Scale | (A) = 25 wm; scale II (B) = 5 um.

38 POSTILLA 161

(p. 299). His massive specimens, given the provisional name C. sordida, pos-
sessing no microscleres, are most probably not clionids.

Tabb and Manning (1961) gave no description of their specimens of C. carib-
baea from the gulf coast of Florida, but stated that de Laubenfels’ papers on the
West Indian Porifera were used to identify sponges found during the course of
their study.

C. caribbaea, again in the gamma stage, was reported from North Carolina
by Wells, Wells and Gray (1960). The specimens, bright orange-yellow in life
and dark brown after preservation, do not possess spirasters. The surface “ex-
hibits many prominent circular areas about 5 mm in diameter. These probably
represent low papillae that have contracted.” (p. 230). These specimens are most
probably not C. caribbaea.

Table | compares the spicular morphology and dimensions, shape and di-
mensions of the excavations, and the color of the type of C. caribbaea, C. viridis
from the Mediterranean, C. viridis from Jamaica (Hechtel’s specimens), and the
specimens collected during the course of the present study.

Carter’s C. caribbaea is doubtless very close to Schmidt’s C. viridis. How-
ever, since differences, as presented above and in Table | do exist, it is deemed
best to retain Carter’s species, at least tentatively, for the widely distributed
West Indian clionid. The relatively recent identifications of C. caribbaea by de
Laubenfels and Wells er a/. are misleading and care should be taken by investi-
gators using these sources as a guide to the identification of West Indian
clionids.

Cliona delitrix, n. sp.

Diagnosis. A very destructive species that always encrusts the surface of its
substrate with a thin layer of red to red-orange tissue. The surface of the sponge,
most frequently elliptical in outline with a maximum diameter of nearly one
meter, is dotted with small white zoanthids. White areas of dead coral surround
the periphery of the sponge. The oscules are large and prominent. The spicula-
tion is composed solely of rather long straight to slightly curved tylostyles, 279
pum by 8.9 wm (mean length by mean width of six specimens). Long cylindrical
galleries, approximately 4 mm in width, extend directly from the encrusting sur-
face to a depth of as much as 10-12 cm within the substrate.

Color. In life, the surface is red to red-orange; the internal portions are ver-
milion. Underwater, the sponge appears brown to the diver. After fixation in
10% neutralized formalin and transfer to 70% ethyl alcohol, it is orange brown.

Dried specimens are cinnamon in color.

Substrates. Diploria labyrinthiformis (Linnaeus), Siderastrea siderea (Ellis &
Solander), Montastrea cavernosa (Linnaeus) and dead coral (massive heads).

JAMAICAN EXCAVATING SPONGES 29

Depths. Collected from 22-34 m. This sponge has not been observed to occur
above 22 m in the study area. It has, however, been collected by the late Prof.
T.F. Goreau at 12 m from Maiden Cay, near Port Royal, Jamaica (YPM No.
4611).

Spicules. (1) Abundant tylostyles (Fig. 7), most with a long, thick shaft tapering
slightly toward the pointed end. A few tylostyles are rather slender (5-6 4m in
width). Most of the tylostyles are straight; some may be slightly curved. Two
categories of heads may be distinguished. A) Heads bearing subterminal knobs
with a maximal diameter of 8.1-10.9 4m. These knobs are quite irregular in
shape. Often the knob is developed on only one side of the head. B) Elongate,
slightly developed heads with a maximum diameter of 7.3-9.8 um. Some of

Fig. 7. Tylostyles of Cliona delitrix n. sp. Scale = 25 um.

30 POS TILEAWGI

these spicules possess a head region so indistinctly separated from the shaft that
they tend toward being styles. Occasionally, there is a swelling behind the head
region near the center of the shaft. Tylostyle length: 279 + 4.6 wm, 205-321
pm. N=6 specimens. Tylostyle width: 8.9 + 0.1 gm, 5.1-10.7 pm. N=6
specimens. Fifty spicules were measured for each specimen. (2) No micro-
scleres were observed.

Surface. This sponge always tends to overgrow the surface of its substrate but
has not been observed to extend up above the substrate as do the gamma stages
of C. celata or C. viridis. Only a thin layer of tissue overlies the surface. In the
early stages, C. delitrix forms discrete ostial and oscular papillae but quickly
begins to overgrow the substrate. When the sponge inhabits large coral heads,
such as Siderastrea siderea or Diploria labyrinthiformis, it is elliptical in outline,
may reach a diameter of nearly one meter (long axis), and occupies the central
region of the coral. White areas of dead coral are seen around the periphery of
the sponge. It appears that this sponge, once established on a coral colony, ex-
pands by actively killing the coral immediately surrounding it. Species of coral
with smaller heads may become completely overgrown and, thus, unrecogniz-
able. The oscular papillae are quite large and the oscules are prominent. Fully
expanded oscules (Fig. 25) may attain a diameter of nearly | cm. There may
either be discrete ostial papillae raised slightly above the surface of the sponge
or the entire surface, with the exception of the oscular papillae, may be covered
with ostia. The latter case is more common in older specimens. Diameter of el-
liptical oscular papillae: 7.85 + 0.14 mm, 7.44-8.21 mm (short axis); 9.65 +
0.10 mm, 8.88-9.92 mm (long axis). N=4 specimens. Diameter of circular ostial
papillae 1.33 + 0.04 mm, 1.26-1.41 mm. N=3 specimens. Diameter of elliptical
ostial papillae: 3.05 + 0.09 mm, 2.79-3.21 mm (short axis); 4.05 + 0.07 mm,
3.89-4.25 mm (long axis). N=4 specimens. The surface of most specimens is
dotted with numerous individuals of the small, white zoanthid, Parazoanthus
parasiticus (Duch. & Mich.) Verrill (Fig. 25). This zoanthid has been previously
reported in association with sponges from Bermuda (Verrill, 1900), the Ba-
hamas (Duerden, 1903), Jamaica (Duerden, 1900, as Parazoanthus separatus),
and the lesser Antilles (Duch. & Mich., 1860, as Zoanthus parasiticus). The
column has a diameter of 2-3 mm; its walls, particularly toward the base, are
impregnated with pieces of debris, sponge spicules and, in this case, calcium
carbonate chips extruded by the host sponge. The density of inclusions in the
column wall of the retracted polyp causes the tissues to be “firm and opaque
white” (Duerden, 1903, p. 500). When, however, the polyps are fully expanded,
these tissues are “quite delicate and transparent to a degree probably not met
with elsewhere among the zoanthids” (p. 500). The short tentacles appear light
brown in color due to the presence of zooxanthellae. Zoanthids are absent from
the smaller, and presumably younger, specimens of this sponge.

Excavations. Long cylindrical galleries, all oriented in a direction perpendicular
to the surlace of the infected coral, may be seen extending directly from the
encrusted surlace downward into the substrate. The sponge completely fills the

galleries it produces but is not confined to them; it also lines those portions of

JAMAICAN EXCAVATING SPONGES 31

the substrate remaining between the galleries with a thin layer of its tissue. Be-
low the oscular papillae, nearly 3 cm of the corallum may be removed. The par-
titions between adjacent galleries of large, older specimens are often removed by
the sponge. When the flesh of the sponge is removed with sodium hypochlorite
solution, the remaining coral skeleton is lacy and very thin. This species ap-
parently excavates the tabular endothecal dissepiments and occupies the central
axis of the corallites constructed by individual polyps. The coral remnants be-
tween corallites are pierced by circular holes, of varying sizes, through which the
lobes of the sponge are interconnected in life. Thus, when coral heads infected
in such a manner are viewed in longitudinal section, the entire skeleton appears
vermilion to a depth of as much as 10-12 cm. The structure of the skeleton is
greatly weakened by the activity of this extremely destructive clionid.

Internal Structure. The abundant tylostyles, scattered throughout the rather
loosely organized tissue, are oriented parallel to the surface in the mesohyl. In
this region, there are large elliptical lacunae (303 um by 101 wm). Granular
golden brown pigment cells are very abundant, particularly in the exopinaco-
derm. These cells (12-15 ,.m in their larger diameter) appear capable of assum-
ing a variety of forms and in this respect are reminiscent of the pigment bearing
“cellules spheruleuses” described by Topsent (1900) for C. schmidti. Choanocyte
chambers are 23-27 xm in diameter.

Holotype. YPM No. 8715. Discovery Bay, Jamaica. Inhabiting Siderastrea
sidera from 37 m. Collected by R.S. Jackson and R. Keeley, Oct. 21, 1968. Re-
positories of paratypical material: BMNH and SUNY-UWI. Eight specimens
from the Discovery Bay reefs were studied.

Comparison With Other Species Of The Genus Cliona. In its habit of over-
growing large areas of the surface of the substrate and of lining the substrate
remaining between adjacent galleries with a thin layer of tissue, C. delitrix is
similar to the Bermudan specimens of C. /ampa. Both exhibit beta but not
gamma stages of development. However, the two have quite different comple-
ments of spicules. Most species of the genus Cliona possess tylostyles with well-
defined globose or subterminal knobs. Many of the tylostyles of C. delitrix have
elongate, slightly developed heads. A few are styles. C. lesueuri (Topsent, 1888,
pp. 80, 88, pl. VII, fig. 8) also has large tylostyles with little or no development
of the head region, but possesses spirasters in addition and does not encrust. The
pattern of the galleries is also different from that of C. delitrix. The removal of
endothecal dissepments of individual polyps, a boring habit characteristic of
C. delitrix, is believed to be unique among the clionids thus far described.

Remarks. The specific name is from the feminine Latin noun delitrix meaning
“a destroyer”.

32 POSTILLA 16!
Cliona peponaca, n. sp.

Diagnosis. This species, exhibiting no tendency to overgrow the substrate,
possesses numerous separate oscular and ostial papillae with the former nearly
twice as large as the latter. The spiculation is composed solely of moderately
long, stout, slightly curved tylostyles, most bearing subterminal knobs. Tylostyle
dimensions are: 243 »m by 3.9 ~m (mean length by mean width of four speci-
mens). Papillae and internal lobes are pumpkin-colored (light orange) in life.
Relatively large, cylindrical galleries, approximately 1.2 cm in diameter, lying
one after another, are interconnected by slender strands of tissue.

Color. In life, the papillae and internal lobes are pumpkin-colored (light orange).
Underwater the sponge appears brown to the diver. After fixation in 10%
neutralized formalin and transfer to 70% ethyl alcohol, it is dark orange-brown.

Substrates. Acropora cervicornis (Lamarck), Porites furcata (Lamarck), and
Agaricia agaricites (Linnaeus) forma agaricites encrusting coral rock.

Depths. Collected from approximately 15 to 29m.

Spicules. (1) The abundant tylostyles (Fig. 8) are moderately long and stout;
most have a slightly curved shaft, but some are straight. The head is most fre-
quently subterminal with a maximum diameter of 8.2-9.6 ym. The knob is
symmetrical and always appears to be equally developed on either side of the
shaft. The tip is blunt and rounded. Occasionally it is quite elongated, and the
knob is set back along the shaft a distance of approximately 10-12 ym. A few
spicules have terminal, globose knobs. The shaft increases slightly and very
gradually in diameter to a point slightly behind the midregion of the spicule. It
then tapers gradually to the pointed end. Tylostyle length: 243 + 2.6 ym, 207-
269 um. N=4 specimens. Tylostyle width: 8.9 + 0.1 wm, 7.2-9.4 wm. Fifty spic-
ules were measured for each specimen considered. (2) No microscleres were
observed to be present.

Surface. Specimens of this sponge exhibited no tendency to undergo papillary
fusion or to overgrow the surface of the substrate. Ostial and oscular papillae
are numerous and always appear to be separate. Ostial papillae are quite small;
the oscular papillae are, on the average, nearly twice as large. The perforations
through which ostial and oscular papillae protrude are circular in outline. The
papillae are fairly abundant but form no distinct pattern over the surface of the
substrate. Oscular papillae bear centrally located oscules that are short and cy-
lindrical, having a diameter of 0.9-1.0 mm when they are fully expanded. Ostial
papillae are low and sievelike. Diameter of ostial perforations: 0.69 + 0.07 mm,
0.49-0.96 mm. N=4 specimens. Diameter of oscular perforations: 1.1 + 0.1 mm,
0.72—1.35 mm. N=4 specimens.

Excavations. The large, cylindrical excavations, completely filled with the soft
tissue of the sponge, are interconnected by slender strands of tissue. When an

JAMAICAN EXCAVATING SPONGES 33

Fig. 8. Tylostyles of Cliona peponaca n. sp. Scale = 25 ym.

infected coral is viewed in cross section, the excavations may be seen to occur
side by side, separated by rather thick (2-3 mm wide) pieces of the coral skele-
ton. In branches of Acropora cervicornis or Porites furcata two galleries are
usually seen in cross section whereas in colonies of Agaricia agaricites forma
agaricites there are often 4-5 consecutive galleries. The galleries in the latter sub-
strate are somewhat elongated compared to those in A. cervicornis or P. furcata.
The lobes of the sponge extend back into their substrate a distance of 0.8-0.9 cm.
Diameter of the galleries in A. cervicornis and P. furcata: 1.2 + 0.2 mm, 0.9-1.5
mm. N=3 specimens. Diameter of the galleries in A. agaricites: 1.4 + 0.4 mm,
1.1-7 mm (long axis); 0.74 + 0.1 mm, 0.69—-0.89 mm (short axis). N=1 specimen.
The channels or tunnels leading from the galleries within the substrate are rather
wide and short.

Holotype. YPM No. 8719. Inhabiting Agaricia agaricites forma agaricites
encrusting coral rock at 27 m, Discovery Bay, Jamaica. Collected by R. Keeley
Pang, Nov. 17, 1969. Repositories of paratypical material: BMNH and SUNY-
UWI. Six specimens from the Discovery Bay reefs were studied.

34 POSTILLA 161

Comparison With Other Species Of The Genus Cliona. Topsent’s (1891) fifth
category of clionids with a spiculation composed solely of tylostyles includes
eight species, all described by Hancock: C. alderi, C. gorgonoides, C. angulata,
C. radiata, C. globulifera, C. quadrata, C. insidiosa, and C. millepunctata.
The first six of these have been placed in synonymy with C. celata (Topsent,
1900). C. insidiosa and C. millepunctata differ from C. peponaca in possessing
smaller tylostyles with globose or elliptical heads, respectively, and excavations
and papillae of relatively small diameter. Hancock’s descriptions of all eight of
the above species (1849, 1867), for the most part based on dried specimens, are
not detailed enough to permit subsequent recognition of his species. In addition,
as mentioned previously, Hancock’s type specimens are apparently no longer in
existence.

In 1915 Annandale, working with the clionids of India, described a new
species, C. kempi, a sponge he believed to be closely related to C. Jobata Han-
cock and C. michelini Topsent, although it only possessed a spiculation of tylo-
styles. The tylostyles, 127-205 um by 4.1-8.2 wm, are rather short and slender,
bear subterminal knobs and taper to very fine points. Papillae and excavations
are said to be minute.

Cliona langae, n. sp.

Diagnosis: A brown-black, zoanthid-bearing sponge that tends to overgrow the
substrate completely, covering it with a thin layer of tissue. The surface is
covered with small ostia interspersed with rather large, prominent oscules. The
internal lobes, dull yellow in color, completely fill the galleries. Spiculation com-
posed of long stout tylostyles, 357 wm by 13.3 wm (mean length by mean width
of five specimens) and spiny spirasters, 31.3 wm by 1.6 wm.

Color. In life, the surface is brown-black, the internal lobes are dull yellow. At
all depths from which it was collected, the sponge appears black to the diver.
After fixation in 10% neutralized formalin and transfer to 70% ethyl alcohol,
the surface is pale grey and the interior is colorless. Dried specimens have a pale
brown surface and pale yellow interior.

Substrates. Montastrea annularis (Ellis & Solander), Acropora cervicornis
(Lamarck), Eusmilia fastigiata (Pallas), overgrowing the surface of a colony,
and dead coral (massive heads).

Depths. Collected from 15-33 m. A specimen of C. langae from 36 m was
photographed by H.M. Reiswig.

') Numerous, long stout tylostyles (Fig. 9A). The shaft is most fre-
d to be straight. Very rarely, it is slightly curved. The diameter
stricts slightly below the head region. Maximum diameter is at

ne spicule, and the shaft tapers very gradually from this region

JAMAICAN EXCAVATING SPONGES

Fig. 9. Spicules of Cliona langae n. sp. A. Tylostyles. B. Spiraster. Scale | = 25 ym;
scale I] = 5 wm.

toward the pointed end. The knob is always terminal and quite globose, with a
maximum diameter of 12.2-16.8 4m. The spicules do not tend to exhibit any
variety in form of the knob or curvature of the shaft. Tylostyle length: 357 +

35

36 POSTILLA 161

3.1 um, 303-407 pm. Tylostyle width: 13.3 + 0.16 wm, 9.0-15.9 wm. (2) Abun-
dant spiny spirasters (Fig. 9B). The low, thin spines spiral around the axis of the
spiraster and are rather widely spaced. They do not branch. Some spirasters
have 2-3 turns; many are almost straight. All are quite slender. Spiraster length:
31.3 + 0.7 pm, 28.3-35.0 wm. N=7 specimens. Fifty spicules in each category
were measured for each specimen.

Surface. The sponge tends to overgrow the surface of its substrate completely,
covering it with a thin layer of tissue. It has not been observed to enter the mas-
sive or gamma stage. C. /angae is most frequently found inhabiting relatively
small (less than one meter in diameter), flattened coral heads of species such as
Montastrea annularis or M. cavernosa on the fore reef slope. The surface of
the sponge is composed of ostial areas interspersed with rather large oscules,
4-6 mm in diameter when fully expanded. The sponge often appears as a cush-
ion-shaped mass of tissue completely enveloping the surface of its substrate.
Most specimens of this species are observed to possess numerous, scattered in-
dividuals of the small, white, epizoic zoanthid Parazoanthus parasiticus
(Duch. & Mich.). This zoanthid has also been observed to occur in association
with C. delitrix and has been described in detail in the section concerning that
species. Two specimens of C. /angae were found inhabiting dead branches of
A. cervicornis. The first specimen, from a depth of 16 meters, formed a small
cushionlike patch, only | cm in diameter, on the surface of the coral. Oscules,
fully expanded were only slightly larger than | mm in diameter. This sponge
possessed a few zoanthids. The second specimen was collected from 33 meters
and is the deepest specimen of the species collected during the course of the
study. This sponge did not encrust or tend to overgrow its substrate. Instead, it
had quite large, separate ostial and oscular papillae. The perforations through
which the papillae protrude are circular. Diameter of ostial perforations:
3.89 + 0.37 mm, 3.46-4.07 mm. Diameter of oscular perforations: 3.94 + 0.39
mm, 3.61-4.12 mm. N=1 specimen. One specimen of this sponge, also bearing.
zoanthids, was found completely encrusting the top of a colony of Eusmilia
fastigiata. This coral bears individual corallites at the ends of branches that are
quite close to one another.

Excavations. The relatively large galleries are irregular in outline and are often
confluent. The excavations extend from the surface of the substrate to a depth
of as much as 3-4 cm within the corallum. In life, the soft, delicate tissue of the
sponge completely fills the excavations and also covers the thin remnants of
coral skeleton remaining between adjacent galleries. When an infected coral
head is viewed in longitudinal section, it may be seen that the network of gal-
leries forms a brown-yellow band immediately beneath the overgrown surface.
[he excavations of the sponge greatly weaken the structure of the coral skeleton.
Diameter of the excavations: 2.31 + 0.09 mm, 2.07-2.78 mm. N=6 specimens.

Internal Structure. The tissue of the sponge is dense and contains tylostyles
strewn in contusion. Tylostyles also form the framework of the oscules. Although
spirasters are present in the choanosome, lining canals, they are most abundant

JAMAICAN EXCAVATING SPONGES 37

in the encrusting surface tissue. Two types of spherular cells are present. Small,
eosinophilic, spherular cells, 6.3-8.2 xm in diameter, are abundant and con-
spicuous throughout the mesohyl and choanosome. The less numerous, larger,
elongated pigment cells (15.7 zm by 5.2 um) bear smaller globules and are dark
brown. This species possesses unicellular, symbiotic algae, 7.4-8.9 ym in di-
ameter, that appear pale yellow-brown in eosin and hematoxylin-stained sec-
tions. Sections stained with toluidine blue reveal a moniliform structure of the
nucleus suggesting that they are zooxanthellae. These algae are quite abundant,
particularly throughout the superficial layers of the sponge. The spherical
choanocyte chambers are 20-24 42m in diameter. One specimen (collected at
Discovery Bay, Jamaica, on Nov. 3, 1969, from 30 m by H.M. Reiswig) con-
tained numerous, large, ovoid eggs, 100 um by 72 um, with a lightly staining
nucleus (28 wm in diameter) and a very darkly staining nucleolus.

Holotype. YPM No. 8716. Inhabiting dead coral from 30 m, Discovery Bay,
Jamaica. Collected by R. Keeley Pang, Oct. 28, 1968. Repositories of paratypi-
cal material: BMNH and SUNY-UWI. Eight specimens from the reefs of
Discovery Bay and Rio Bueno, Jamaica, were studied.

Comparison With Other Species Of The Genus Cliona. Species of the genus
Cliona possessing long, stout tylostyles and long, rather thin, spiny spirasters
include C. celata, C. viridis, C. subulata, C. caribbaea, and C. orientalis Thiele
(1900). Of these, C. swbulata and C. orientalis were considered by Vosmaer (1933)
to be insufficiently described. C. caribbaea and its relation to C. viridis are dis-
cussed above. C. celata and C. viridis both become massive, passing through a
very transitory beta stage. Although C. /Jangae does not appear to be close to
either of these sponges, it is interesting to note that C. viridis from the Mediter-
ranean also possesses symbiotic algae (Sara and Liaci, 1964; Sara, 1965). C.
aprica, a new species from Jamaica, with relatively long, stout tylostyles and
spiny spirasters, is described below. It differs from the present species in the de-
gree of encrustation of the substrate, the oscular dimensions, spicular dimen-
sions and spination of the spirasters, as well as the dimensions and form of the
excavations and internal portions.

Remarks. The species name ts given in honor of Dr. Judith C. Lang, who first
discovered the presence of this, and several other, species of clionids on the
Jamaican reefs.

Cliona laticavicola, n. sp.

Diagnosis. Sponge with dull orange papillae and red-orange internal lobes,
excavating very wide galleries that are completely filled with its tissue, and hav-
ing a spiculation composed solely of tylostyles. Two forms occur. One, from
shallow water (0-5 m), possesses scattered ostial and oscular papillae that often
undergo fusion, and long, stout, slightly curved tylostyles, 349 4m by 10.2 um

38 POSTILLA 161

(mean length by mean width of six specimens); the other, from deeper water (12-
17 meters), possesses smaller papillae that do not tend to undergo fusion, and
has shorter, thinner tylostyles, 279 wm by 7.3 wm. Tylostyles of both forms most
frequently bear subterminal knobs.

Two forms of this species occur on the reefs of Discovery Bay, one from
shallow water and the other from deeper water. Each of the forms is described
separately below.

Cliona laticavicola forma laticavicola

Color. In life, the papillae are orange, the internal lobes are red-orange. Very
shallow specimens (1-2 meters) will appear orange to the diver whereas speci-
mens from slightly deeper water (3-5 meters) appear orange—brown. After fixa-
tion in 10% neutralized formalin and transfer to 70% ethyl alcohol the sponge is
dark orange-brown. Dried specimens are pale orange-brown.

Substrates. Diploria labyrinthiformis (Linnaeus), Montastrea annularis (Ellis
& Solander), Porites porites (Pallas), P. astreoides (Lesueur) encrusting old
coral rock, Acropora palmata (Lamarck), dead coral, and coral rock.

Depths. Specimens from 0-3 m were collected and preserved. The sponge was
observed to occur commonly in A. pa/mata to a depth of 5 m. Specimens from
5 m were collected and examined in the laboratory but not preserved.

Spicules. (1) Long, relatively stout tylostyles (Fig. 10), most possessing a
straight shaft, some with the shaft slightly curved. The greatest diameter of the
shaft is in the midregion of the spicule. The shaft tapers quite abruptly from a
point approximately 15 «4m above the pointed end of the spicule. The knob is
most frequently observed to be subterminal with a maximum diameter of 12.2-
14.0 um. The tip is very blunt and rounded. The head region of the spicule may
exhibit a rather wide variety of shapes. They are as follows: (a) head elongated,
somewhat spatulate, with a maximum diameter of 10.8-12.1 wm, (b) head with
an asymmetrically subterminal knob more developed on one side than the
other, (c) knob subterminal but with the swelling set back away from the tip a
distance of 5-8 um, (d) head region devoid of any swelling or differentiation
from the shaft, spicule possesses a slight swelling near the central region and
looks rather like a style. Spicules in this category are rare. Ditylote spicules are
not numerous but a few are present in each of the above categories. Tylostyle
length: 349 + 3.1 um, 256-404 um. Tylostyle width: 10.2 + 0.1 wm, 6.7-12.6 wm.
N=6 specimens. Fifty spicules were measured for each specimen considered. (2)
No microscleres were observed to be present.
the scattered separate ostial and oscular papillae are very numerous,
iite large. Although they form no distinct pattern over the surface of
often undergo fusion. The perforations through which the
rude are quite circular, those of the larger papillae are

JAMAICAN EXCAVATING SPONGES 39

Fig. 10. Tylostyles of Cliona laticavicola n. sp. forma laticavicola. Scale = 25 pm.

rather irregular in their outline. Even the larger, and presumably older, speci-
mens have not been observed to overgrow the surface of their substrate. The
oscules are short (approximately 3 mm in height), wide, and rounded, having
their greatest diameter at the base. When fully expanded, the diameter of the
oscular opening is | mm or less. The largest oscular papillae have excentrically

40 . POSTILLA 161

positioned oscules and also exhibit ostial areas. This is most probably due to
papillary fusion. The ostial papillae are sievelike and form low mounds lying
close to the surface of the substrate. Diameter of ostial perforations: 2.55 + 0.13
mm, 2.09-3.31 mm. N=7 specimens. Diameter of oscular perforations: 2.83 +
0.17 mm, 2.41-3.78 mm. N=7.

Excavations. Large galleries completely filled with the rather tough tissue of the
sponge (Fig. 23). This species inhabits a wide variety of substrates. The shape
and dimensions of its galleries are, to a certain extent, dependent upon the shape
and dimensions of the substrate. One of the most common substrates, Acropora
palmata, forms large, flattened, frondlike branches. In pieces of this coral hav-
ing a thickness of approximately | cm, the sponge forms elongated flattened
galleries that are somewhat sausagelike in their appearance. Adjacent galleries
are separated by thin remnants of the substrate (1-2 mm in width). Diameter of
the galleries in A. pa/mata: 16.0 + 1.4 mm, 9-20 mm (long axis); 6.0 + 0.89
mm, 4.4-7.9 mm (short axis). N=4 specimens. The galleries leave only 3-4 mm of
the coral skeleton remaining at either surface of the coral. In the rather stout
branches of colonies of Porites porites, the sponge forms cylindrical excavations.
When an infected branch is viewed in cross section, the sponge may be seen as a
mass of tissue occupying the central axis. In outline, it forms a circle with a very
regular circumference. Diameter of the galleries in P. porites: 14.0 + 0.09 mm,
8.3-17.2 mm. N=4 specimens. A single lobe of the sponge may extend along the
axis of the coral for 2-3 cm. Subsequent lobes are interconnected by very slen-
der (1-2 mm in diameter) stems of various lengths. When the sponge inhabits
the more massive coral heads (Diploria labyrinthiformis, Montastrea annularis)
or massive pieces of old coral rock, it forms very extensive galleries with an ir-
regular or ragged outline. Diameter of the galleries in massive substrates: 4.6 +
0.27 cm, 3.8-6.0 cm (long axis); 3.2 + 0.17 cm, 2.2-4.8 cm (short axis). N=4
specimens. In such substrates the galleries lie rather near one of the surfaces. In
all substrates the channels leading from the internal lobes of the sponge to the
surface openings are quite short and wide. Length: 4.4 + 0.15 mm, 3.8-5.7 mm.
Width: 2.8 + 0.07 mm, 1.7—3.4 mm. N=7 specimens.

Internal Structure. The tylostyles of the internal lobes are strewn in confusion.
Choanocyte chambers are 21-24 wm in diameter. Two types of spherular cells
are present. Golden brown pigment cells appear capable of assuming a variety
of shapes, from spherical to elongate. They are rather small, 6.3-8.9 jm in di-
ameter. The second type of spherular cell, localized in the choanosome, are
elongated and more than twice as large as the pigment cells (14-18 pm by 5-7
uum). They have highly eosinophilic granules, a pale, large nucleus and promi-
nent nucleolus.

Notes On C. laticayvicola forma laticavicola From Puerto Rico. The Porifera
collection of the Peabody Museum at Yale contains a specimen (YPM No.
‘414B) of C. laticavicola forma laticavicola from 1-3 m of water at Cayo de
Caballo Ahogao, near La Parguera, Puerto Rico. The sponge, collected in 1959,

y Dr. W.D. Hartman, was found inhabiting old, dead pieces of Acropora pal-

JAMAICAN EXCAVATING SPONGES 4]

mata and the central axis of a living colony of Millepora sp. It appears to be
exceptionally vigorous in the extent of its growth. This specimen closely re-
sembles those Jamaican specimens described above in regard to spicular mor-
phology, surface morphology, and the extent and configuration of the excava-
tions. Measurements made on the preserved specimen reveal the following
statistics: tylostyle length, 339 + 4.4 4m (mean + standard deviation), 301-363
pm (range); tylostyle width, 8.9 + 0.2 pm, 7.3-10.6 um (fifty spicules were
measured); diameter of ostial perforations, 3.27 + 0.11 mm, 2.81—3.39 mm; di-
ameter of oscular perforations, 3.41 + 0.16 mm, 2.99-4.00 mm; diameter of
excavations, 1.8 + 0.1 cm, 1.1-2.4 cm.

Cliona laticavicola forma parvispiculata

Color. In life, after preservation or drying, the color is as described for C. Jati-
cavicola forma laticavicola. The sponge appears brown to the diver.

Substrates. Acropora cervicornis (Lamarck). The sponge has not been observed
to occur in any other substrates.

Depths. Collected from 12-20 m.

Spicules. (1) Tylostyles like those of C. laticavicola form laticavicola but shorter
and somewhat thinner (Fig. 11). The spicules most frequently have subterminal
heads and exhibit variations in the shape and position of the knob that are
similar to those described above for C. laticavicola forma laticavicola. Tylostyle
length: 274 + 2.8 wm, 248-324 um. Tylostyle width: 7.3 + 0.18 ym, 5.0-9.2 pm.
Fifty spicules were measured for each specimen considered. (2) No microscleres
were observed.

Surface. The numerous ostial and oscular papillae are similar in their mor-
phology to those of the shallow water form described above. They are, however,
slightly smaller. Diameter of ostial perforations: 2.18 + 0.05 mm, 1.15—3.26 mm.
N=4 specimens. Diameter of oscular perforations: 2.54 + 0.03 mm, 1.60—3.87
mm. N=3 specimens. The sponge does not overgrow the surface of its substrate,
but papillary fusion may be observed.

Internal Structure. As described for the shallow water form of the species.

Holotype. The holotype of C. /aticavicola is YPM No. 8720. This specimen,
representative of the shallow water form of the species, was found inhabiting
coral rock from 3 m. Collected by R. Keeley Pang and P.K.T. Pang, November
29, 1969, at Discovery Bay, Jamaica. Six specimens from the reefs of Discovery
Bay were studied. (Paratypical material includes YPM No. 8721, representative
of the deep water form. Inhabiting A. cervicornis from 12 m, Discovery Bay,
Juamaica). Repositories of other paratypical material: BMNH and SUNY-UWI.

A? POSTILLA 161

Fig. 11. Tylostyles of Cliona laticavicola n. sp. forma parvaspiculata. Scale = 25 pm.

Comparison With Other Species Of The Genus Cliona. C. laticavicola may be
distinguished from all other species of clionids by its exceptionally wide, cav-
ernous galleries and spiculation composed solely of large, stout tylostyles, most
frequently having asymmetrical, subterminal knobs.

Remarks. The species name is from the Latin /atus (wide) + cavea (excavation)
+ -colus (inhabiting, dwelling).

Cliona aprica, n. sp.

sponge, when young, possesses numerous small, separate
sstial and oscular papillae. Although older specimens exhibit

JAMAICAN EXCAVATING SPONGES 43

extensive papillary fusion, the sponge never completely overgrows the surface of
the substrate. The internal lobes, dull yellow in life, completely fill the spherical
to ellipsoidal galleries. Two forms of this species occur. One, from shallow water
(0-5 m), with a spiculation composed of tylostyles, 326 »m by 8.4 um (mean
length by mean width of six specimens) and spiny spirasters, 34.1 »m by 1.9 wm,
has larger, stouter oscules and overgrows the substrate more extensively; the
other, from deeper water (12-34 m), possesses smaller, thinner tylostyles, 286 ~m
by 7.9 wm and slightly smaller spirasters, 33.9 zm by 1.8 zm. Each of the forms
is described separately below.

Cliona aprica forma aprica

Color. In life, the surface is brown-black, the interior is yellow. At all depths
from which it was collected, the sponge appears black to the diver. After fixation
in 10% neutralized formalin and transfer to 70% ethyl alcohol, the surface is
pale grey, the interior is colorless. Dried specimens have a very pale brown sur-
face and pale yellow interior.

Substrates. Acropora palmata (Lamarck), Porites astreoides (Lesueur), dead
coral, and coral rock.

Depths. Collected from 0-5 m.

Spicules. (1) Abundant tylostyles (Fig. 12A) with a relatively stout shaft that is
most frequently slightly curved but is sometimes straight. The shaft has its
greatest diameter near the midregion and tapers very gradually toward the
pointed end. The knob, with a maximum diameter of 9.3-11.4 ym, is always
terminal. It is globose to elliptical; the globose forms predominate. Frequently
there is a secondary annular swelling behind the head region. This swelling may
occur immediately behind the knob or may be some 5-10 um behind it. The
spicules do not exhibit great variety in their form. Tylostyle length: 326 + 3.1
pm, 284-356 zm. N=6 specimens. Tylostyle width: 8.4 + 0.16 ym, 7.8-10.6 wm.
N=6 specimens. (2) Spirasters (Fig. 12B) are common but not abundant. They
are located primarily within the surface tissue and may be overlooked in spicule
preparations made only with tissue from the galleries. The spirasters may be
spiral with 2-3 turns, C-shaped or almost straight. All possess spines, 1-2 .m in
length, along the axis and at either end of the axis. Although the spines are
always observed to be restricted to the convex side of the axis, their form,
number and arrangement are variable. Length of spiral spirasters: 34.1 + 1.4 wm,
29.2-39.8 zm. Width of spiral spirasters: 1.9 + 0.07 wm, 1.42.5 wm. N=7 speci-
mens. Length of straight spirasters: 17.9 + 0.18 um, 16.1-19.4 wm. Width of
straight spirasters: 7.4 + 0.10 um, 6.9-7.8 «um. Length of C-shaped spirasters:
14.3 + 0.17 pm, 12.2-15.9 wm. Width of C-shaped spirasters: 2.6 + 0.05 pm,
2.0-3.7 ym. Fifty spicules from each of the two categories were measured for
each specimen considered.

44 POSTIEE AVG!

Fig. 12. Spicules of Cliona aprica n. sp. forma aprica. A. Tylostyles. B. Spirasters.
Scale | (A) = 25 wm; scale I] (B) = 5 wm.

Surface. The sponge always tends to overgrow the surface of its substrate. Very
young specimens have numerous, separate ostial and oscular papillae that pro-
trude through nearly circular perforations in the substrate. The papillae are ir-
regularly distributed over the surface, forming no distinct pattern. Oscules (Fig.
21) are small, stout and have their greatest diameter at the base. Viewed from
above with a dissecting microscope, the ostial papillae present a reticulated pat-
tern. Ostial papillae are raised up above the surface of the substrate. The num-
erous ostia are confined to the top surface of the papillae and are approximately
15-20 «xm in diameter. Fusion of the papillae is quite common, even in young

JAMAICAN EXCAVATING SPONGES 45

specimens. As the sponge begins to overgrow the substrate, the oscules become
surrounded by ostial areas until almost all the papillae are confluent. Tiny (7—-
10 «m in diameter) chips of calcium carbonate may often be seen surrounding
the base of the oscules. Almost all of the specimens of C. aprica forma aprica
observed or collected encrusted a large portion of their substrate with a thin
layer of tissue. Diameter of circular ostial perforations: 0.52 + 0.06 mm, 0.30-
0.64 mm. N=4 specimens. Diameter of elliptical perforations of ostial papillae
and of compound papillae (ostial and oscular): 1.05 + 0.07 mm, 0.80-1.84 mm
(short diameter); 1.67 + 0.09 mm, 0.96-3.20 mm (long diameter). N=6 specimens.

Excavations. The relatively small spherical to ellipsoidal galleries are quite ir-
regular in outline and very frequently are confluent. The soft tissue of the
sponge completely fills, and is always confined to, the galleries. In substrates
such as Acropora palmata the excavations lie close to the overgrown surface
and only extend approximately 8-9 mm down into the skeleton of the coral.
Channels leading from the galleries to the surface papillae are very short (1-2
mm) and relatively wide (2-3 mm in diameter). Diameter of galleries: 1.20 +
0.09 mm, 1.08—1.89 mm. N=6 specimens.

Internal Structures. The tylostyles are abundant in the surface layers, with the
pointed ends directed outward. Sections through the ostial papillae reveal a
band of parallel tylostyles comprising the framework of the papillae. Spirasters,
most abundant in the surface layers at the base of papillae, are also present in
the choanosome. The tylostyles of the internal lobes are strewn in confusion.
Two types of spherular cells are present. The highly eosinophilic spherular cells
have a diameter of 8.4-10.5 4m. The brown pigment cells are larger, with a di-
ameter of 11.7-14.0 xm, but have smaller globules. The species possesses uni-
cellular, symbiotic algae (8.1-10.0 ym). Spherical choanocyte chambers are
18-22 wm in diameter.

Cliona aprica forma profunda

Color. As for C. aprica forma aprica. The surface, in life, is slightly paler,
sometimes almost grey.

Substrates. Acropora cervicornis (Lamarck), Solenastrea bournonii Milne
Edwards & Haime, Agaricia agaricites (Linnaeus) forma agaricites, A. agari-
cites forma purpurea, Montastrea annularis (Ellis & Solander), M. cavernosa
(Linnaeus), Porites furcata (Lamarck), Lucina pensylvanica (bivalve), dead
coral and coral rock.

Spicules. (1)Abundant tylostyles (Fig. 13A) resembling those of Cliona aprica
forma aprica in their form, but with the shaft slender rather than stout and con-
siderably shorter. The last one-third of the shaft of a few tylostyles is rather
sharply bent. Tylostyle length: 286 + 2.3 pm, 276-297 ym. Tylostyle width: 7.9
+ 0.11 um, 7.7-8.2 wm. N=9 specimens. (2) Spirasters (Fig. 13B) have the same

46 POSTILLA 161

Fig. 13. Spicules of Cliona aprica n. sp. forma profunda. A. Tylostyle. B. Spiraster.
Scale | = 20 ym; scale Il = 5 um.

location and form as those described tor C. aprica forma aprica. They are
slightly smaller. Fifty spicules from each of the two categories were measured
tor each specimen considered.
Surface, As described for the shallow water form, this sponge always tends to

surface of the substrate. The progression from separate ostial and |
‘ae through papillary fusion to the encrusting of the substrate is |

JAMAICAN EXCAVATING SPONGES 47

also seen in this form. However, the encrusting of the surface has not been ob-
served to proceed as far as that of the shallow water form. The dimensions
of the papillary perforations are similar to, but slightly smaller than, those given
for C. aprica forma aprica. When fully expanded, the oscules are short cylinders
of uniform diameter. They are slightly smaller than the oscules of C. aprica
forma aprica.

Excavations. As described for the shallow water form the galleries are irregular
in outline and often are confluent. When the sponge inhabits massive substrates
(M. annularis, M. cavernosa, S. hyades), its excavations are confined to the
superficial portions of the corallum. However, in the rather slender branches of
A. cervicornis, C.-aprica forma profunda may produce a network of galleries
extending from one surface of the branch to the opposite one.

Internal Structure. As described for C. aprica forma aprica. However, both the
eosinophilic spherular cells and the algae are less numerous in the deep water
form.

Holotype. The holotype of C. aprica is YPM No. 8722. This specimen, repre-
sentative of the shallow-water form of the species, was found inhabiting A.
palmata from 2 m, Discovery Bay, Jamaica. Collected by R. Keeley, March 28,
1969. Six specimens from the reefs of Discovery Bay were studied. (Paratypical
material includes YPM No. 8723, representative of the deep water form of this
species. Inhabiting A. cervicornis from 15 m, Discovery Bay, Jamaica.) Reposi-
tories of other paratypical material are: BMNH and SUNY-U WI.

Comparison With Other Species Of The Genus Cliona. C. aprica is compared
with C. /angae in the discussion of the latter species.

Family SPIRASTRELLIDAE

Anthosigmella varians (Duch. & Mich.) de Laubenfels

Thalysias varians Duchassaing and Michelotti, 1864, p. 86.
Anthosigmella varians, de Laubenfels, 1936a, p. 143.

(non) Anthosigmella varians, Wells, Wells and Gray, 1960, p. 55.
Suberites tuberculosus Schmidt, 1870, p. 46.

Suberites coronarius Carter, 1882, p. 352.

(non) Suberites coronarius, Carter, 1897, p. 74.

Spirastrella coronaria, Topsent, 1894, p. 26.

Anthosigmella coronarius, Topsent, 1918, p. 557.

(non) Cliona coronaria, Dendy, 1916, p. 132.

Papillina arcuata Topsent, 1889, p. 35.

48 POSTILLA 161

Color. In life, the surface is brownish tan, the interior is dull yellow. After fixa-
tion in 10% neutralized formalin and transfer to 70% ethyl alcohol, the sponge is
beige. Dried specimens are pale brown.

Substrates. Acropora cervicornis (Lamarck) and dead coral (massive heads).
Depths. Collected from 20-41 m.

Material. Four specimens from the reefs of Discovery Bay, Jamaica, were
studied.

Spicules. (1) Abundant, long, rather slender tylostyles (Fig. 14A), tapering
gradually toward the pointed end. Most have a slightly curved shaft, a few are
straight. Occasionally the shaft may be arched or bent at the midregion. There
is considerable variation in the shape of the head region: (a) head elongated,
well developed with a constriction in the neck region; (b) head elliptical with
small annular swelling immediately behind it; (c) knob subterminal, tip rounded
and blunt; (d) development of head much reduced; spicules are stylelike. These
are rare. Tylostyle length: 423 + 2.9 um, 271-465 pm. Tylostyle width: 8.7 +
0.12 um, 7.9-10.4 wm. N=4 specimens. (2) Abundant anthosigmas (Fig. 14B);
most are arched and C-shaped, a few are almost straight. There are prominent,
knoblike spines on the convex side and at each extremity. The knobs at the
extremities are frequently branched and those along the axis may also be
branched. Anthosigma length: 19.4 + 0.14 pm, 15.4-24.9 um. Anthosigma
width: 2.2 + 0.17, 1.9-2.5 wm. N=4 specimens. Fifty spicules in each category
were measured for each specimen considered.

Surface. Sponge completely encrusting the surface of the substrate with a thick
layer of rather rubbery tissue. Ostia are minute. Oscules are quite small and
inconspicuous, approximately 2~3 mm in diameter.

Interior. Internal portions of sponge inhabit rather large hollows or cavities in
the upper portion of the corallum of dead coral. The cavities resemble the ex-
cavations produced by C. aprica. Evidence is inconclusive as to whether A.
varians is capable of producing the cavities through its own activity or whether
it merely infiltrates spaces already present within the coral skeleton. Diameter of
cavities: 2.9 + 0.07 mm, 2.2-3.8 mm. N=3 specimens.

Internal Structure. As described by Hechtel (1965, p. 56). Anthosigmas are most
abundant in the interior of the sponge, particularly in the choanosome where
they are found lining canals. Symbiotic algae, 8.9-10.3 wm in diameter, are
present but not abundant.

Taxonomic Discussion. Despite the presence of highly distinctive microscleres,
the taxonomic history of A. varians has been quite complex. The original de-
scription (as Thalysias varians) by Duchassaing and Michelotti in 1864, although
insufficient, emphasized the great variation in shape that this species may un-
dergo. The spiculation of the sponge was neither described nor figured. Schmidt

JAMAICAN EXCAVATING SPONGES 49

3

|

Fig. 14. Spicules of Anthosigmella varians (Duch. & Mich.). A. Tylostyles. B. Antho-
sigma. Scale 1 (A) = 25 wm; scale 11 (B) = 5 um.

(1870) believed that he was dealing with a new species, Suberites tuberculosus,
that took many forms: encrusting, massive or branching. The presence of spiras-
ters was not mentioned. Carter (1882) proposed the new species, Suberites coro-
narius, for a West Indian sponge with peculiar spirasters. In 1889 Topsent

50 POSTHECARION

described Papillina arcuata, with arched spirasters bearing knoblike, branched
spines, from the Campeche Bank. Then, in 1894, Topsent placed P. arcuata in
synonomy with Suberites coronarius Carter and transferred Carters species to
the genus Spirastrella. In 1913 he established a new genus, Anthosigmella, for
sponges with the distinctive anthosigmas as microscleres, and transferred
Spirastrella coronaria (Carter) Topsent to that genus. Topsent (1920) later re-
examined Schmidt’s type specimen and, after finding anthosigmas, placed it in
synonomy with A. coronarius. However, in 1936, de Laubenfels transferred
Tha lysias varians Duch. & Mich. to the genus Anthosigmella and placed Carter's
species in synonomy with it.

Geographical Distribution. West Indies: Jamaica, Carter, 1882, p. 352 (as
Suberites coronarius), Hechtel, 1965, p. 55; Bahamas, Carter, 1882, p. 352 (as
Suberites coronarius), de Laubenfels, 1949, p. 19; Honduras, Carter, 1882, p.
352 (as Suberites coronarius), Lesser Antilles, Duchassaing and Michelotti, 1864,
p. 86 (as Thalysias varians from Guadeloupe, St. Barthelemy, St. Thomas and
Tortole). Florida: Schmidt, 1870, p. 46 (as Suberites tuberculosus). Gulf of
Mexico: Campeche Bank, Topsent, 1889, p. 35 (as Papillina arcuata); Florida
Gulf coast, de Laubenfels, 1953, p. 539, Little, 1963, p. 55.

A. varians, under a variety of names, has often been reported from the West
Indies, Florida and the Gulf of Mexico. It is apparently endemic to this region.

Discussion: The specimens of A. varians from Discovery Bay were all of the
encrusting form with the internal portions of the sponge inhabiting cavities
within corals. None of the previous reports of this species have mentioned this
habit. De Laubenfels wrote of A. varians, “This common West Indian sponge is
strikingly marked by its occurrence in coral sand with a buried substratum
from which digitate processes arise” (p. 203).

However, Carter (1887) in a record of the presence of his species Suberites
coronarius from the Mergui Archipelago, stated that the lower portion of the
sponge is “mingled with a layer of coral which has been cancellated by the
excavating habit of the sponge” (p. 74). Thiele (1900), who believed that the
Mergui sponge was not the same as the West Indian sponge, redescribed the
former as Cliona orientalis from specimens from Ternate, Indonesia. Annandale
(1915) re-examined a portion of Carter’s material from Mergui and concluded
that Thiele had been correct in his assumptions. Dendy (1916) reported Suber-
ites coronarius dwelling in cavities in corals. Based on this, he transferred the
species to the genus Cliona. Although the specimens reported by Carter (1887)
and Dendy (1916) are most probably clionids and not A. varians, it should be
noted that A. varians does occupy an extensive network of cavities in corals.

(Order and Family Uncertain)
Alectona jamaicensis, n. sp.

Diagnosis. Sponge with numerous, small, separate, pale-yellow ostial and os-
cular papillae and relatively large spherical to ellipsoidal galleries. Spiculation

JAMAICAN EXCAVATING SPONGES 51

composed of the following three categories, the first two exhibiting a wide range
of variation in form: (1) stout diactinal spicules, 215 pm by 10.8 wm, (mean
length by mean width of one specimen), covered with rounded tubercles that
may be either low or prominent but are never branched or spined; (2) thin,
smaller diactinal spicules, 203 zm by 7.9 ym; (3) amphiasters, 59 jum by 2.2 wm,
bearing a whor] of 5-6 tylote rays near each extremity.

Color. In life, pale yellow. Underwater, the sponge appears to be almost color-
less to the diver. After fixation in 10% neutralized formalin and transfer to 70%
ethyl alcohol, it is completely colorless.

Substrate. Porites furcata Lamarck.

Depth. Collected from 14 m. Only one specimen was collected during the course
of the present study.

Spicules. The spiculation of the genus Alectona is highly distinctive. (1) Rela-
tively few, stout diactinal spicules (Fig. 15B) of moderate length, curved or
rather sharply bent in the midregion of the axis, and exhibiting a wide range of
variation in their forms. Some are recurved at one end; some form a V with un-
equal arms. The diameter of the diactinal spicules is slightly greater in the cen-
tral region than at either end. The spicules may either taper uniformly or possess
a swelling or protuberance along the convex side of the axis, often near the
center. The extremities are obtuse rather than pointed. The spicules are covered
with rounded tubercles, apparently not arranged in any distinct pattern. The
tubercles of an individual spicule may be low (less that | wm in height) or quite
prominent (3-4 ym in height), numerous or rather sparsely distributed, but the
rounded extremities of the spicule always bear many low tubercles over their
surface. The tubercles are never branched or spined. These spicules, all quite
unusual in their appearance, are of a type not seen elsewhere within the Hadro-
merida. Length: 215 + 2.4 wm, 186-237 wm. Width: 10.8 + 1.2 wm, 9.6-12.3 um.
Twenty-five spicules were measured in this category. (2) Numerous thinner,
slightly smaller diactinal spicules (Fig. 15A), usually smooth but sometimes
bearing scattered flattened tubercles. Shaft tapers from the midregion to pointed
extremities at either end. These spicules, as those described above, exhibit a
wide range of forms. (a) The axis may be bent rather sharply at the center and
possess a swelling in this region, on the convex side giving the spicule a winged
appearance. (b) The axis may be recurved at one end. (c) The central swelling
may appear as a knot or twist in the spicule. (d) The axis may be nearly straight
and bear a spherical swelling at its center. Length: 203 + 1.7 wm, 186-287 um.
Width: 7.9 + 1.2 «wm, 6.1-8.5 um. Fifty spicules were measured in this category.
The swelling or protuberance is reported to be the remnant of a third ray (Top-
sent, 1900). See Discussion. Some spicules of the present specimen possess a
rudimentary second ray, giving them a clublike or tylostylelike appearance.
Length: 112 + 1.1 pm, 97-144 ym. Twenty-five spicules were measured in this
category. (3) Abundant amphiasters (Fig. 15C) of rather precisely determined
form and a wide range of lengths. The axis is of a uniform diameter, smooth
except for its pointed, microspined extremities. Near each extremity is a whorl

52 POSTILLA 161

Fig. 15. Spicules of Alectona jamaicensis n. sp. A. Thin diactinal spicule. B. Thick
diactinal spicule. C. Amphiaster. Scale | (A,B) = 25 wm; scale 11 (C) = 5 wm.

of rays, usually 5-6 in number, projecting outward from the axis, each ray bear-
ing small spines terminally. Some rays have a slightly greater diameter at their
most distal, microspined end. Length: 59 + 1.4 wm, 31-74 wm. Width: 2.2 +
0.08 zm, 1.4—4.3 jum. Fifty spicules were measured in this category.

Surface. Ostial and oscular papillae are separate, rather small, and low, lying
close to the surface of the substrate. Fusion of the ostial papillae was not ob-
served. The perforations through which the ostial and oscular papillae protrude
are circular in outline. This specimen exhibited no tendency to overgrow the
substrate. Ostial papillae are sievelike in appearance, bearing numerous small
ostia. The oscules are shaped like truncated cones. Diameter of the ostial per-
forations: 0.51 + 0.06 mm, 0.39-0.72 mm. N=8 measurements. Diameter of the
oscular perforations: 0.54 + 0.08 mm, 0.41-0.83 mm. N=6 measurements.

JAMAICAN EXCAVATING SPONGES 33

Excavations. Relatively large galleries, roughly spherical to ellipsoidal in their
outline. Subsequent lobes are interconnected by cylindrical stems approxi-
mately 2 mm in diameter. Diameter of the galleries: 6.2 mm (mean), 5.8—7.9 mm
(range). Three galleries were measured. The single specimen collected was
found inhabiting the dead, encrusted basal portion of a relatively stout branch
of Porites furcata (diameter: 1.2 cm). In terms of the quantity of substrate re-
moved by the sponge, A. jamaicensis is a rather destructive species of boring
sponge. See Discussion. The galleries are completely filled with masses of the
soft sponge tissue. The internal lobes of the sponge communicate with the sur-
face papillae through short, quite narrow channels. Diameter of the channels:
0.42 mm, 0.37-0.51 mm. Length of the channels: 3.2 mm, 2.8-3.9 mm. N=7
measurements.

Holotype. YPM No. 8718. Discovery Bay, Jamaica. Inhabiting Porites furcata,
from 14 m. Collected by R. Keeley Pang, Nov. 10, 1969.

Taxonomic History Of The Genus Alectona. The genus Alectona (from Alecto,
one of the furies, Carter, 1879c, p. 494) was established by Carter in 1879 for
excavating sponges possessing the following spiculation: “Skeleton-spicule
acerate, abruptly curved or bent in the center, tubercled throughout. Flesh-
spicule spindle-like, consisting of a straight shaft, pointed at the extremities and
encircled by two rings of tubercles equidistant from each other and from the
ends of the shaft respectively.” The genus includes: A. wallichi Carter, 1874, p.
252; A. millari Carter, 1879c, p. 494; A. higgini Carter, 1880, p. 58; and A. prim-
itiva Topsent, 1933, p. 554. A. wallichi was originally described as Gummina
Wallichii. The description was based on an isolated spicule found in an arena-
ceous deposit from the Agulhas Shoal, Cape of Good Hope. This spicule,
according to Carter, resembled very closely an isolated spicule figured by
Bowerbank (1864, p. 270, pl. xi, fig. 244) and described as having been re-
peatedly found in the matter obtained by washing the roots of Oculina rosea
(Stylaster sanguineus fide Carter, 1879b, p. 353). In 1879 Carter redescribed G.
Wallichii from two deep water specimens from the collection of the British
Museum (Natural History). Due to an error in editing, it appeared as Corticium
Wallichii (1879b, p. 353). Later on in the same year, the description of the new,
closely allied species, A. millari, led Carter (1879c, p. 496) to transfer G. wal-
lichii to the new genus A/ectona and to consider it as an excavating sponge. A.
millari has received no names in synonomy. Carter (1879c) suggested that it
might be a variety of A. wallichi but subsequent investigators have considered
it as a separate species. A. higgini was found excavating melobesian nodules in
the Gulf of Manaar. Its flesh spicules or amphiasters are quite similar to those
of A. wallichi and A. millari. However, it also possesses two other distinctive
categories of spicules: (1) rather stout diactinal spicules with rounded extrem-
ities and “a number of annular depressions and inflations, the latter of which
are microspined” and (2) “fine hair-like acerate” spicules “tending toward the
form of a tricurvate” (Carter, 1880, p. 58). De Laubenfels (1936) believed that

54 POSTILLA 161

this species was not of the genus A/ectona and, perhaps, not even a sponge. He
would establish the new genus Delectona for it. Earlier, Topsent (1891) had,
however, noted the similarities between the spiculation of A. higgini and Dotona
pulchella Carter, 1880, p. 57 (also originally described from melobesian nodules
of the Gulf of Manaar) and suggested that the latter species might well be trans-
ferred to the genus A/ectona or that if the genus Dotona were to be conserved
A. higgini should become Dotona higgini.

Geographical Distribution Of The Genus Alectona. |. Alectona millari. At-
lantic: between the North of Scotland and the Faroe Islands, Carter, 1879c, p.
494: coast of Norway, Topsent, 1900, p. 27; Azores, Topsent, 1900, p. 25.
Mediterranean: Banyuls, Topsent, 1900, p. 25; Gulf of Naples, Sara, 1959, p.
10, Russ and Riitzler, 1959, p. 767 (as Alectona sp.), Riitzler, 1966, p. 313. Red
Sea: coast of Sudan, Topsent, 1920, p. 555. Il. Alectona wallichi. Indian Ocean:
Agulhas Shoal, Cape of Good Hope, Carter, 1874, p. 252 (as Gummina wallichi,
from an isolated spicule); Seychelles, Carter, 1879b, p. 353 (as Corticium wal-
lichi). “South Seas”: Carter, 1879b, p. 353 (as Corticium wallichi). Il. Alectona
higgini. Indian Ocean: Gulf of Manaar, Carter, 1880, p. 58. IV. Alectona
primitiva. Indian Ocean: coast of South Australia, Topsent, 1933, p. 555.

Comparison With Other Species Of The Genus Al/ectona And Discussion. The
present species is very close to Topsent’s specimens of A. millari, particularly
that from the Mediterranean at Banyuls. It was recognized by Topsent (1933)
that the specimens he had described as A. millari in his 1900 monograph actually
differed somewhat in spicule morphology and dimensions from the type de-
scribed by Carter in 1879. Unfortunately, since Carter’s and Topsent’s speci-
mens were obtained from dredgings, little is known of the surface morphology
of this species in life.

Until recently A. millari was known only as a deep-water species (from depths
of 450 m or more). Volz (1939) did not find A. millari at Rovigno, and regarded
it as being solely from deep water (p. 72). However, Sara (1959) mentioned the
presence of A. millari in the shallow waters of the Grotto of Gaiola (Gulf of
Naples, Tyrrhenian Sea) co-occurring with Cliona copiosa Sara, C. celata, C.
viridis, and Cliothosa hancocki (Schmidt) Topsent. Russ and Riitzler (1959),
also working in the Gulf of Naples, recorded the presence of several specimens
of a yellow-white boring sponge, Alectona sp. In the report of a subsequent
study of that area, Riitzler (1966) stated that the specimens referred to as
Alectona sp. by Russ and Riitzler (1959) belong to A. millari. He found addi-
tional specimens boring in the dead bases of Petrobiona massiliana Vacelet and
Sara (Order Pharetronida, Family Murrayonidae). Unfortunately, neither Sara

nor Rutzler provided a description or figures of their specimens of A. millari. It
is interesting to note that all the shallow-water Mediterranean specimens of this
ponge were taken from shaded localities or underwater caves.

Isolated spicules, presumed to belong to A. millari, were found by Sara (1959)
in tne sponges Topsentia contorta Sara and Haliclona viscosa Sara, from the

JAMAICAN EXCAVATING SPONGES a5

shallow waters of the Adriatic. Riitzler (1965) also reported the presence of
isolated spicules of A. millari in the sponge Terpios fugax Duch. & Mich. col-
lected from a depth of one meter in the North Adriatic. Neither of these two
reports included figures of the spicules. However, Riitzler described two cate-
gories of spicules from his specimens: robust diactinal spicules bearing tuber-
cles, 120-220 um in length; and amphiasters with tylote rays, 15-18 jm in
length. It appears impossible at this time to judge whether these investigators
were dealing with A. millari as originally described by Carter, A. millari sensu
Topsent (1900), the form found in the shallow waters (14 meters) of Jamaica,
or with something rather different from all three of the above.

Previous collections of boring sponges from the Discovery Bay reefs, made
by the late Prof. T.F. Goreau and E. A. Graham in 1962 (deposited in the Pea-
body Museum of Natural History, Yale University) and by Dr. J.C. Lang in
1966-67 did not reveal the presence of this species of Alectona. Only one speci-
men was observed during the course of the present study. It is most regrettable
that other Jamaican or, indeed, West Indian specimens are not available for
comparison.

Topsent noted irregularities in the spiny diactinal spicules of his specimen
from Banyuls; he considered these to be individual teratological variations.
However, much the same irregularities in form were seen in the spicules of the
Jamaican specimen. Topsent did not consider the smooth, smaller diactinal
spicules as belonging to a separate category. He stated that they are merely
poorly developed forms of the larger, stouter diactinal spicules and noted that
in both of these kinds of spicules the axial canal turns outward the swelling or
protuberance. He took this to mean that these spicules are derived from a
three-rayed form and that the swelling is the vestige of this third ray. Indeed,
Carter (1879), who also noted the turning of the axial canal, figured and de-
scribed three-, four- and five-rayed spicules. These spicules have not been
reported from subsequent specimens of A. mi//ari and have not been found in
the Jamaican specimen. A second category of spicules found only by Carter is
composed of spicules that are “acerate undulating, almost immeasurably fine,
hair-like, with an enlargement in the centre barrel-shaped, inflated in the
middle and at the ends respectively” (p. 495). These were not figured.

The diactinal spicules, figured and described by Carter (1879) are considerably
stouter (28 zm) than those from the Jamaican specimen (9.6-12.3 um, range).
The tubercles, according to both Carter and Topsent are conical and may be
branched at the extremities or may bear spines. Due to the differences in the
classes of spicules present, their morphology and dimensions and to the vast
differences in habitats (the very deep, cold waters of the North Atlantic as op-
posed to the shallow, warm waters of Jamaica), the present specimen is placed
in the new species, A. jamaicensis.

56 POSTILLA 161

Order HAPLOSCLERIDA

Family ADOCIIDAE
Siphonodictyon brevitubulatum, n. sp.

Diagnosis. Sponge forming a single large, spherical excavation completely filled
with soft, pale-yellow, mucus-covered tissue. Lemon yellow ostial and oscular
papillae are separate. The hill-like ostial papillae, composed solely of a network
of numerous small ostia, are confluent in some specimens. Spiculation of
smooth, rather abruptly pointed oxeas, 134 wm by 8.0 wm (mean length by mean
width of four specimens).

Color. In life, papillae are lemon yellow; the mass of sponge within the exca-
vation is pale, dull yellow. Underwater, the sponge appears yellow to the diver.
After fixation on 10% neutralized formalin and transfer to 70% ethyl alcohol the
papillae are pale yellow, and the internal portions are drab yellow-tan.

Substrates. Porites furcata Lamarck, Solenastrea bournonii Milne Edwards &
Haime, Montastrea annularis (Ellis & Solander), M. cavernosa (Linnaeus),
and Acropora cervicornis (Lamarck).

Depths. Collected from 5—32 m.

Spicules. (1) The abundant, smooth oxeas (Fig. 16) are relatively stout. They
may be curved, sharply bent at the midpoint of the axis, or almost straight. The
oxeas do not taper but are rather abruptly pointed. Neither styles nor strongyles
have been observed. There is not great variety in the shape of the spicules or in
the curvature or termination of the axis. Oxea length: 134 + 2.7 wm, 119-148
pm. Oxea width: 8.0 + 0.2 wm, 6.9-9.1 »m. N=4 specimens. Fifty spicules were
measured for each specimen considered. (2) No other categories of spicules were
observed to be present.

Surface. The sponge does not tend to overgrow the substrate. Oscular and
ostial papillae (Fig. 24) are separate and relatively few in number. Oscules,
protruding through circular perforations in the substrate, have not been ob-
served to be surrounded by ostial areas. The oscules are cylindrical and rather
prominent (7-9 mm in height) in comparison to those of most clionids. The
perforations through which the ostial papillae protrude are elliptical. In some
specimens ostial papillae are discrete and widely separated, while in others sev-
eral papillae are confluent. These hill-like papillae, comparable in height to the
oscules, are composed solely of a network of numerous small ostia. Diameter
of oscular perforations: 3.11 + 0.14 mm, 1.98-3.72 mm. N=4 specimens. Diam-
eter of ostial papillae: 1.71 + 0.25 mm, 0.98-2.04 mm (short axis); 2.03 + 0.31
mm, 1.82~2.79 mm (long axis). N=4 specimens. Confluent ostial papillae are
approximately 7-8 mm by 2 mm (long axis by short axis).

JAMAICAN EXCAVATING SPONGES 57

Fig. 16. Oxeas of Siphonodictyon brevitubulatum n. sp. Scale = 20 pm.

Excavations. The activity of the sponge produces a single, large spherical exca-
vation completely filled with the soft, mucus-covered tissue of the sponge. In
massive substrates (S. bournonii, M. annularis and M. cavernosa) the exca-
vation may attain a diameter of 4-5 cm. The diameter of the excavations in
branching substrates (P. porites and A. cervicornis) is considerably less, 1—2
cm. The tunnels or channels leading from the excavated cavity in the substrate
to the surface openings are narrow and elongated. Channel length in massive
substrates: 8.1 + 0.1 mm, 7.2-9.8 mm. N=4 specimens. Channel length in
branching substrates: 4.2 mm, 3.8-5.9 mm (two specimens). Since S. brevitubu-
latum most frequently inhabits very massive coral heads, it is difficult to bring
intact specimens of this species to the laboratory for study. However, when the
specimen infecting A. cervicornis was examined in the laboratory, it was ob-

58 POSTILLA 161

served actively expelling tiny calcium carbonate chips through the excurrent
openings.

Holotype. YPM No. 8717. Inhabiting Montastrea annularis at 15 meters, Dis-
covery Bay, Jamaica. Collected by R. Keeley, Oct. 16, 1968. Repositories of
paratypical material: BMNH and SUNY-UWI. Seven specimens from the reefs
of Discovery Bay, Jamaica, were studied.

Comparison With Other Species Of The Genus Siphonodictyon. The genus
Siphonodictyon, closely related to the genus Phloeodictyon Carter, was estab-
lished by Bergquist (1965) for mucus-producing sponges of cryptic habit, lacking
spongin and a barklike dermal region, and possessing a dermal skeleton com-
posed of brushes of oxeas. The type species of the genus, Siphonodictyon mu-
cosum Bergquist, may be distinguished from the present species by its color
(black in life), the brittle texture and length of the tubes (2.0-7.4 cm) termin-
ating in oscules or sieve areas, as well as the dimensions of the oxeas (194-212
pm by 8.5-11.5 4m). Bergquist’s species is from 3-20 ft. in the Palau Islands.
Riitzler (1971) has described two new species of the genus Siphonodictyon (S.
cachacrouense and S. coralliphagum) from the West Indies. He distinguished
four forms of S. coralliphagum: forma typica, forma obruta, forma tubulosa
and forma incrustans. Of these, S. coralliphagum forma tubulosa from Scotts
Head Bay, Dominica, W. I., is closest to S. brevitubulatum. However, S. coral-
liphagum forma tubulosa forms dense clusters of ostial and oscular tubules with
the oscular tubules attaining a height of more than twice that of the ostial tu-
bules. Concerning the oxeas, Riitzler stated that “the tips are rather blunt to
mammiform, usually smooth, sometimes rough to stepped. Some styles and
strongyles occur” (p. 9, and fig. 7). In addition, the oxeas, which are said to
occur in two size categories, are longer but thinner than those of the present
species. The spicular dimension of Riitzler’ss USNM #24101 (specimen repre-
sentative of the form), based on 100 measurements, are: 156.0—-180.8 by 5.0-7.1
um (large oxeas) and 121.6-174.4 by 1.3-4.0 »m (small oxeas).

Remarks. The species name is a compound of the Latin adjectives brevis

(short) and tubulatus (tubular), for the distinctive short oscular tubes of the Ja-
maican specimens.

IV. Key to the Excavating Sponges Described from Discovery Bay, Jamaica

ulation including tylostyles ..c:...2,.11-.cst esses ssooteos ase ese ee 2
on not including tylostyles. 2 c.ci05-.<.5-<--s2o0ccdsoes eee o eee 12
les, NO MICTOSClEFES: <...02555.020g.se0sccsedeosaeessernece ace eee

JAMAICAN EXCAVATING SPONGES 59

3. Encrusting; surface red to red-orange; often with small white
zoanthids; tylostyles, 205-321 by 5.1—-10.7 pm .................... Cliona delitrix
I NUGNA MEIN CHUIS (NN) Steere ee eee ere ass aoe ole eases suds dacs dvoabins ndulawceeaeioeedakdeechoncs -

4. Pointed end of tylostyles attenuated or “ragged”; tylostyles,

134-221 by 7.8-11.4 um; tiny (0.60-0.70 mm) dull yellow
GstialeamMa osc ulate papillae y.cs...c6<hsakgeecsss:ssseedvossocewssedeecestesteasanes C. janitrix
Rompediendioftylostyles motattemuated): .. 22. ccieccicsivescoseseceos ceconecteecvenceosoeona OD

5. No papillary fusion; tylostyles, 207-269 by 7.2-9.4 1m,
pumpkin-colored; oscular papillae almost twice as
large as ostial papillae................ Uasrbensiersaesnes ee VCD ONACG
Papillary fusion; red to Peon cle pallericg
tylostyles, 256-404 by 6.7-12.6 um; depth
GM CMO OM etre neuen se akt bees acta ees esta saan. xsieGeioigs doen alls stuaeas savas C. laticavicola
forma /aticavicola

tylostyles, 248-324 by 5.0-9.2 um; depth

GANDER) SENN ae en eee Pac <P eso oc tenict node ewneeuesubteseadesads C. laticavicola
forma parvispiculata

6. Tylostyles, oxeas and microrhabds; tylostyles few in
number, 236-288 by 4.0-7.1 um; microspined oxeas,
60-116 by 2.1—4.3 um; spiny microrhabds, 9.222. 1
by 1.1-2.6 yxm; vermilion; oscular papillae (0.25—
3.44 mm) larger than ostial papillae (0.18—1.48 mm) .................. C. lampa
MESTOSta ME STAINES DUE ASUE IS Mercere tae) 20 00 aaccsic chan Piet seacclenent csvveladaesscavegnassiteseeds setae 7

7. Spirasters smooth, wormlike, multiply bent, 31.8—52.7
by 2.2—4.7 xm; tylostyles, 141-387 by 5.0-10.2 um;
red to red-orange; papillary perforations,
Oe OSI A Srnec sake scio voanaie deets suse aeceeusbleneres C. vermifera
RIAU AS UTS ESO MMC Ue acee treme sine SEs aera a ciseesas see edodt ocean ceticy soolns uw oteuiessevescenss teks 8

8. Spirasters are anthosigmas, usually C-shaped, with
prominent, branched spines; size of anthosigmas,
15.4-24.9 by 1.9-2.5 pm; tylostyles, 389-477
by 7.9-9.2 4m; brownish tan; encrusting ............... Anthosigmella varians
SS PIEASLCRSMEUILGIT PIN CMIL ede essed sac besa ste ce aicdeecsatwns<u dtu costes desecacunisetuenaveceee 9

9. Spirasters of two types: short and stout, 26.5-68.9
by 4.1-7.9 um; long and thin, 60.9-100.7
by 1.6-3.2 wm. Striking purple coloration
RE MIAUMIS TAM CT PLESCUVALION® 0.272.5..cceac--aceseetonnssu ssa 2shavaaseuaes acces C. schmidti
PSUS UC IS (ONO ETL IC eee ss cnn ops adeac nr vencev odes encresneiecenttaaise(sherasassnestyeeeaae 10

10. Completely overgrowing substrate, usually with
small, white zoanthids; sponge black—brown;
tylostyles, 303-407 by 9.0-15.9 wm; spirasters,
Des 35) OMDV IED US IM, Sop esiaen «con wanreaveviancsale «szentnesseneeetiezanwcdver soe C. langae
MNOUCOMIPICLE IVIOWET PLOWS SUDSELALC: .........20<.c-sscantnear'ssdatssrecaesiesteeeesceeaerer 1]

60 POSTILLA 161

11. Extensive papillary fusion; brown-black; tylostyles,
284-356 by 7.8-10.6 wm; spirasters, 29.2-39.8
by 1.4-2.5 wm; depth range, 0-5 Mm ......5...0..s:2000-00essiseoossnenasoreeeee eel OC mMGPUC
forma aprica
tylostyles, 276-297 by 7.7-8.2 wm; depth
Pane, L234 a0 ce csnvderogudesiecdownanines cebetdeecce eeepeadde sot ese Recetas C aprica
forma profunda
No papillary fusion; dull yellow; tylostyles, 361-451
by 9.4-12.1 zm; spirasters, 27.3-41.8 by
DQ 252 pr 22. ung. sessaecboveuia stan ats <2 cao coven Meauesyeesgn-Senese dot eee eee C. caribbaea

12. Spiculation of smooth oxeas, 119-148 by
6.9-9.1 wm; lemon yellow; oscules, short
prominent cylinders, 7-8 mm in height; ostial
papillae often confluent, diameter
78. Tandy 2 MM =; ee sennessceredeueeeee eet Siphonodictyon brevitubulatum
Spiculation of diactinal spicules and amphiasters;
stout tubercled diactinals, 151-354 by 9.6—12.3
ym; thinner, usually smooth diactinals, 186—
287 by 6.1-8.5 wm; amphiasters, bearing
whorl of 5-6 rays near each extremity, 31-
74 by 1.4-4.3 xm; pale yellow; tiny (0.39-
O:83:mm) papillae: 2.2...cscezeees saeco tee eee Alectona jamaicensis

V. Diversity and Zoogeography of Reef-dwelling Excavating Sponges

The diversity of reef-dwelling clionids as previously reported in the literature
is surprisingly low. A study by Burton (1934) of the sponges collected during
the Great Barrier Reef Expedition revealed the presence of two coral-dwelling
spirastrellids, Spirastrella inconstans (Dendy) and Spirastrella aurivillii Lind-
gren, but no clionids. De Laubenfels (1954) found five species of clionids in the
west central Pacific. The West Indian shallow water sponge fauna is both di-
verse and colorful in most localities. Yet of the 233 species from this region
listed by de Laubenfels (1950b) only two, both endemic, are clionids. One
coral-dwelling spirastrellid, Spirastrella dioryssa de Laubenfels, was also found
to be present. Studies of the sponges of the Lesser Antilles at Curagao (Arndt,
1927) and Barbados (Hechtel, 1969) list no clionids.

From relatively recent investigations of the Porifera of the Bahamas, Ber-
muda (de Laubenfels, 1949, 1950a) and Jamaica (Hechtel, 1965) four species of
clionids are reported: Cliona lampa (Bermuda), C. vastifica (Bahamas), C.
viridis (= C. caribbaea, Jamaica) and C. vermifera (Jamaica). These workers
also reported the presence of Anthosigmella varians from the Bahamas (de
Laubenfels, 1949) and Jamaica (Hechtel, 1965). The low diversity of previously
reported excavating sponges from the West Indies may be contrasted to the
presence of nine species (three endemic) of boring sponges in the shallow waters
of the Adriatic (Volz, 1939).

JAMAICAN EXCAVATING SPONGES 61

Thirteen species of excavating sponges from the reefs of Discovery Bay and
Rio Bueno, Jamaica, are described in the present report (Table 2.) In addition,
eight undescribed species of excavating sponges, not reported in this paper,
have been tentatively identified from the study area. The low diversity or ab-
sence of reef-dwelling clionids in the reports of other workers is most probably
due in large part to the cryptic habit of these sponges. Most excavating sponges
communicate with the external environment by means of small, rather incon-
spicuous incurrent and excurrent papillae. The largest of these papillae may
attain a diameter of 2-3 mm, but quite frequently a species will have papillae
of less than 0.5 mm in diameter.

Of the six previously described species found during the course of the present
investigation, three, C. caribbaea, C. lampa and A. varians, are apparently con-
fined in their distribution to tropical Atlantic America. C. janitrix has only been
described from Corsica previously. C. schmidti and C. vermifera have been
reported most frequently from the Mediterranean and Adriatic seas, but they
also occur in the Indo-Pacific. One of the seven new species, C. laticavicola, is
also found in Puerto Rico. Alectona jamaicensis is very close to Topsent’s
(1900) deep-water Mediterranean specimen of A. millari. The description of
the new species of Alectona from Discovery Bay, Jamaica, represents the first
report of this genus from the waters of the Western Hemisphere. It is also in-
teresting to note that C. caribbaea is in many respects similar to C. viridis,
originally described from the Adriatic by Schmidt (1862)

The faunal affinities of the Jamaican excavating sponges are apparently with
those of the Mediterranean and Adriatic seas. De Laubenfels (1950b) stated
that the faunal affinities of the West Indian Porifera in general are with the
Porifera of the Mediterranean and the East Indies. According to Hechtel (1965)
most of the 57 known species of sponges from Jamaica have a limited distribu-
tion. Two-thirds are endemic to tropical Atlantic America. He speculated that
there might be a strong West African influence in the sponge fauna of the
Lesser Antilles. However, a subsequent study of the Demospongiae of Bar-
bados (1969) gave “no evidence of affinities with West African sponges” (p. 5).
Most similarities are with Indo-Pacific sponges. Very little is known concern-
ing the clionids of the Lesser Antilles.

Competition for space and the protective advantages offered by the indwell-
ing habitus have apparently led to the acquisition of this mode of existence by
several different groups of sponges, particularly within the coral reef environ-
ment. In this environment we find not only numerous species of clionids, but
in addition, coral-dwelling spirastrellids and excavating species of the genus
Siphonodictyon.

VI. Transplant Studies

Two of the new species described above, Cliona aprica and Cliona laticavicola,
were found to have a deep-water and a shallow-water form that may be dis-

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64 POSTILLA 161

tinguished by differences in the dimensions of the spicules and papillae, the
degree of papillary fusion and, to a slight degree, spicule morphology. Trans-
plant studies were conducted with these species in an effort to determine the
influence of the habitat upon the characteristics of the two forms.

Method. Transplants of excavating sponges were made following the method
of Neumann (1966). The substrates used for the transplants were: calcite, Ver-
mont marble, dead shells of the oyster, Crassostrea virginica (Gmelin), and
uninfected branches of Acropora cervicornis and A. palmata that had been
washed in fresh water to remove the living tissue and placed in running sea
water overnight. Small pieces of sponge-infected coral taken from shallow (0-3
m), intermediate (8-16 m) or deep (20-30 m) water were broken open, at-
tached to the various substrates and placed at two stations, one at 15 m ona
coral island (Kinzie, 1970) in a sand channel of the cervicornis zone and the
other at 24 m on a large coral-covered rock situated on the sand between the
cervicornis zone and the fore reef slope. (The terminology of the reef zones
follows that of Kinzie, 1970). The transplants at the 15-m site suffered consid-
erable damage during the winter storms; unfortunately, most were lost during
this period. All transplants at the 24-m site were recovered. The dates of the
transplants were as follows: at the 15-m site, 16 Nov. 1969 to 25 March 1970;
at the 24-m site, 25 March 1969 to 9 Nov. 1969 and 12 Nov. 1969 to 25 March
1970. The mean and range of tylostyle measurements were calculated. Fifty
measurements were made for all specimens considered.

Results. The results of the transplants, discussed below, are summarized in
Tables 3 and 4.

a) Cliona aprica. Small pieces of the shallow- and deep-water forms of this
sponge were transplanted to 15 and 24 m. All but one of the transplants at 15
m were lost during the winter storms. C. aprica forma aprica from A. palmata
at | m was successfully transplanted to both A. palmata and Vermont marble
placed at 24 m. Transplants of both forms of this species on calcite and Cras-
sostrea virginica failed. After 7.5 months, transplants infecting A. palmata and
Vermont marble showed very little tendency to encrust. The original shallow-
water sponge was observed to be growing over a large portion of the substrate
when it was originally collected. The tylostyles of the transplants, although
abundant, were quite thin and much smaller. These new spicules resembled
those of C. aprica forma profunda in appearance. The spicular dimensions of
the original sponge had not changed. However, the color of the papillae had
changed.

Iransplants of C. aprica forma profunda originally inhabiting A. cervicornis
at 18 m to uninfected branches of A. cervicornis at 24 m were also successful.
When examined 7.5 months later, the transplants exhibited extensive papillary

fusion. The compound papillae were slightly smaller than those of the original
sponge. Tylostyle dimensions of the original remained the same in each case.
lylostyles of the transplants were thinner but of a comparable length. The ex-

cavations of the transplants extended through the superficial layers of the

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JAMAICAN EXCAVATING SPONGES 67

branches of A. cervicornis for a length of 7.1-8.3 cm. One transplant of the
deep-water form of C. aprica to Vermont marble placed at the 15-m site sur-
vived the winter storms. Spicular dimensions of the transplant were comparable
to those of the original sponge, although the tylostyles were slightly thinner.

b) C. laticavicola. When small pieces of the shallow-water form of this spe-
cies, originally inhabiting P. porites at 1 m , were transplanted to uninfected
branches of A. palmata placed at 24 m, small elliptical excavations, 2-3 mm by
1 mm (long axis by short axis; three specimens), were produced in the new sub-
strate. The excavations in the original substrate were irregular and circular with
a diameter of 1.0-1.3 cm. In the new substrate, the perforations through which
the papillae protrude were considerably smaller than those of the original
sponge. Papillary fusion, extensive in the original sponge, was not observed to
occur in the transplant. The tylostyle dimensions of the original sponge re-
mained unchanged. The tylostyles of the transplants were very much shorter and
thinner than those of the original.

Small pieces of C. /aticavicola forma parvispiculata inhabiting A. cervicornis
from 18 m were attached to uninfected branches of A. cervicornis and placed
at 24 m for 7.5 months. The transplants excavated a series of cylindrical gal-
leries, 2-3 mm in diameter, extending throughout the central axis of the coral
for a length of 2.0-3.5 cm. The excavations of the original, also cylindrical,
were |.1 cm in diameter and filled the central axis of the infected branch for a
length of approximately 10 cm. The tylostyles of the transplant were compar-
able in length to those of the original, but were thinner. Papillary perforations
of the transplants were also smaller than those of the original sponge. Trans-
plants of the shallow-water and deep-water forms of C. laticavicola to Cras-
sostrea virginica and calcite failed.

Previous investigators, dealing with seasonal variation among natural popu-
lations or with gemmules grown under laboratory conditions, have also found
that environment parameters may affect spicular dimensions. Early experi-
mental work by Jgrgensen (1944, 1947), using the fresh-water sponge Spongilla
lacustris (Linnaeus), showed that an increase in the silica concentration of the
medium resulted in an increase in the uptake of SiO? by the forming spicules.
He reported (1944) that thicker microscleres were formed in a medium con-
taining more silica. The length, however, was not affected.

Hartman (1958) found that in natural populations of the marine sponge
Haliclona canaliculata Hartman winter colonies possess thicker spicules than
summer colonies, and suggested that this might be due to the greater avail-
ability of silicates in New England waters throughout the winter months. Stone
(1970) was able to correlate seasonal changes in the spicules of Hymeniacidon
perleve (Montagu) with the changing levels of environmental silicates in British
coastal waters.

Recent laboratory studies conducted by Elvin (1971) using gemmules of
Ephydatia muelleri (Lieberkiihn), a fresh-water sponge, are in agreement with
those of Jgrgensen, and show that the rate of silica deposition is increased when
the silicic acid concentration of the medium is increased. Although the rate of
growth in spicule length decreases with an increase in silica concentration, the

68 POSTILLA 161

projected final length of spicules in media of higher silica content would be
longer than those in lower concentrations.

It is suggested that the differences in spicular dimensions observed in the
shallow-water and deep-water forms of Cliona aprica and C._ laticavicola
may be due to differences in the concentration of available silicates in the en-
vironments inhabited by these forms.

VII. Acknowledgements

I would like to thank Dr. Willard D. Hartman for his continuing interest and
encouragement throughout this investigation and for his critical reading of the
manuscript.

The late Professor Thomas F. Goreau first gave me the opportunity to study
the excavating sponges of Jamaica at the State University of New York-
University of the West Indies Marine Laboratory. I am deeply appreciative of
his help, encouragement and kindness.

Grateful thanks are given to all those associated with the SUNY-UWI
Marine Laboratory during the time of my field studies. I am particularly in-
debted to the late R. Shean Jackson, laboratory manager and diver.

Discussions with colleagues, Dr. J.C. Lang and Dr. H.M. Reiswig, were of
great value. Underwater photographs were generously donated by Dr. H.M.
Reiswig.

I am grateful to Miss S.M. Stone of the British Museum (Natural History)
for permission to examine specimens of clionids deposited in that institution
and for her kind hospitality.

Mr. Carl Wester and Mrs. Virginia Simon have assisted in the preparation
of the drawings.

My husband, Peter, assisted in part of the field work and with the photog-
raphy. For all his help and advice I am very grateful.

This work was supported in part by a grant-in-aid of research from The
Society of the Sigma Xi.

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rt aa re 1664 A Monograph of the British Spongidae. Vol. 1: 290 p. Ray Soc.,

ion

faurice. 1934. Sponges. Sci. Rpt. Great Barrier Reef Exped., 1928-29. 4: 513-

JAMAICAN EXCAVATING SPONGES 69

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—— — 1953b. A guide to the sponges of Eastern North America. Special Publ. U. of
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JAMAICAN EXCAVATING SPONGES 13

Fig. 17. Oscule of Cliona lampa de Laubenfels inhabiting dead coral. Scale = 2 mm.

Fig. 18. Excavations of Cliona lampa inhabiting dead coral. Scale = 5 mm. Fig. 19.
Excavations of Cliona vermifera Hancock. Scale = 3 mm. Fig. 20. Ostial and oscular
(lowermost edge) papillae of Cliona vermifera Hancock. Scale = 3 mm. Fig. 21. Os-
cule of Cliona aprica n. sp. forma aprica inhabiting dead coral. Scale = 3 mm.

POSTILLA 161

74

JAMAICAN EXCAVATING SPONGES 75

Fig. 22. Excavations of Cliona lampa de Laubenfels in Acropora cervicornis. Scale in
mm. Fig. 23. Excavations of Cliona laticavicola n. sp. forma J/aticavicola in Diploria. Ca.
0.5X. Fig. 24. (right) Oscule and discrete and confluent ostial papillae of Siphonodictyon
brevitubulatum n. sp inhabiting Montastrea. The papillae are surrounded by a red en-
crusting sponge. Photograph by H.M. Reiswig. Scale = 3 mm. Fig. 25. Oscules of Cliona
delitrix n. sp. Epizoic zoanthids, Parazoanthus parasiticus (Duch. & Mich.), may be
seen growing in the sponge surface. Photograph by H.M. Reiswig. Scale = | cm.

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