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Bulletin 20 - Peabody Museum of Natural History - Yale University

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PEABODY MUSEUM OF NATURAL HISTORY<br />

YALE UNIVERSITY<br />

BULLETIN <strong>20</strong><br />

A Systematic Study <strong>of</strong> the Demospongiae<br />

<strong>of</strong> Port Royal, Jamaica<br />

BY<br />

GEORGE JOHN HECHTEL<br />

Department <strong>of</strong> Biology<br />

State <strong>University</strong> <strong>of</strong> New York<br />

Stony Brook, L. I., N. Y.<br />

NEW HAVEN, CONNECTICUT<br />

1965


<strong>Bulletin</strong>s published by the <strong>Peabody</strong> <strong>Museum</strong> <strong>of</strong> <strong>Natural</strong> <strong>History</strong> at <strong>Yale</strong> <strong>University</strong><br />

embody research carried out under the auspices <strong>of</strong> the <strong>Museum</strong>. The issues are numbered<br />

consecutively as independent monographs and appear at irregular intervals. Shorter papers<br />

are published at frequent intervals in the <strong>Peabody</strong> <strong>Museum</strong> Postilla Series.<br />

EDITORIAL BOARD: Elwyn L. Simons, Chairman<br />

Charles L. Remington<br />

N. Philip Ashmole<br />

EDITOR: Ellen T. Drake<br />

Communications concerning purchase or exchange <strong>of</strong> publications should be addressed<br />

to the Editor, <strong>Peabody</strong> <strong>Museum</strong> <strong>of</strong> <strong>Natural</strong> <strong>History</strong>, <strong>Yale</strong> <strong>University</strong>, New Haven, Connecticut<br />

065<strong>20</strong>, U.S.A.<br />

Issued March 1, 1965


A SYSTEMATIC STUDY OF THE DEMOSPONGIAE<br />

OF PORT ROYAL, JAMAICA


Printed in the United States <strong>of</strong> America


LIST OF ILLUSTRATIONS<br />

ABSTRACT<br />

INTRODUCTION<br />

ACKNOWLEDGMENTS<br />

METHODS<br />

HABITATS<br />

PREVIOUS JAMAICAN RECORDS<br />

CLASSIFICATION<br />

TAXONOMIC CRITERIA<br />

SYSTEMATICS<br />

FAMILY Spongiidae<br />

Dysideidae<br />

Aplysillidae<br />

Haliclonidae<br />

Desmacidonidae<br />

Adociidae<br />

Callyspongiidae<br />

Tedaniidae<br />

Microcionidae<br />

Mycalidae<br />

Halichondriidae<br />

Spirastrellidae<br />

Suberitidae<br />

Clionidae<br />

Placospongiidae<br />

Tethyidae<br />

Geodiidae<br />

Chondrillidae<br />

Plakinidae<br />

DISTRIBUTION BY HABITAT<br />

FIELD KEY<br />

ZOOGEOGRAPHICAL NOTES<br />

SUMMARY<br />

LITERATURE CITED<br />

INDEX<br />

PLATES I-VIII<br />

CONTENTS<br />

V<br />

vi<br />

1<br />

2<br />

2<br />

2<br />

3<br />

4<br />

4<br />

5<br />

5<br />

8<br />

14<br />

17<br />

18<br />

21<br />

26<br />

31<br />

37<br />

41<br />

45<br />

52<br />

54<br />

59<br />

60<br />

62<br />

65<br />

68<br />

74<br />

76<br />

77<br />

80<br />

83<br />

84<br />

85<br />

90<br />

95


ILLUSTRATIONS<br />

TEXT-FIGURES<br />

1. Spicules <strong>of</strong> Haliclona hogarthi, p. <strong>20</strong>.<br />

2. Spicules <strong>of</strong> Adocia implexiformis, p. 27.<br />

3. Spicules <strong>of</strong> Adocia albifragilis, p. 28.<br />

4. Spicules <strong>of</strong> Pellina coeliformis, p. 29.<br />

5. Spicules <strong>of</strong> Sigmadocia caerulea, p. 30.<br />

6. Spicules <strong>of</strong> Callyspongia pallida, p. 36.<br />

7. Spicules <strong>of</strong> Microciona microchela, p. 41.<br />

8. Spicules <strong>of</strong> Microciona rarispinosa, p. 43.<br />

9. Spicules <strong>of</strong> Thalyseurypon conulosa, p. 44.<br />

10. Spicules <strong>of</strong> Ulosa hispida, p. 51.<br />

11. Spicules <strong>of</strong> Halichondria magniconulosa, p. 53.<br />

12. Spicules <strong>of</strong> Diplastrella megastellata, p. 58.<br />

13. Spicules <strong>of</strong> Geodia papyracea, p. 71.<br />

14. Spicules <strong>of</strong> Erylus ministrongylus, p. 73.<br />

15. Spicules <strong>of</strong> Corticium tetralophum, p. 77.<br />

PLATES<br />

I.<br />

II.<br />

III.<br />

IV.<br />

V.<br />

VI.<br />

VII.<br />

VIII.<br />

Fig. 1.<br />

Figs. 2,3.<br />

Fig. 4,<br />

Fig. 5<br />

Fig.<br />

Fig. 2<br />

Fig. 3,<br />

Fig. 4<br />

Fig. 1<br />

Figs. 2,3<br />

Fig. 4,<br />

Fig. 5<br />

Fig.<br />

Fig. 2<br />

Fig. 1<br />

Fig. 2,<br />

Fig. 3<br />

Fig. 4<br />

Fig.<br />

Fig. 2,<br />

Fig. 1<br />

Fig. 2<br />

Fig. 3<br />

Figs. 4,5.<br />

Figs. 1,2.<br />

Fig. 3.<br />

Verongia fistularis (Pallas).<br />

Haliclona erina de Laubenfels.<br />

Haliclona doria de Laubenfels.<br />

Oligoceras hemorrhages de Laubenfels.<br />

Haliclona hogarthi n. sp.<br />

Adocia implexiformis n. sp.<br />

Gelliodes areolata (Wilson).<br />

Desmapsamma anchorata (Carter).<br />

Pellina coeliformis n. sp.<br />

Adocia carbonaria (Lamarck).<br />

Sigmadocia caerulea n. sp.<br />

Callyspongia pallida n. sp.<br />

Callyspongia plicifera (Lamarck).<br />

Callyspongia fallax Duchassaing and Michelotti.<br />

Halichondria magniconulosa n. sp.<br />

Halichondria melanadocia de Laubenfels.<br />

Zygomycale parishii (Bowerbank).<br />

Tedania ignis (Duchassaing and Michelotti).<br />

Thalyseurypon conulosa n. sp.<br />

Mycale laevis (Carter).<br />

Placospongia carinata (Bowerbank).<br />

Diplastrella megastellata n. sp.<br />

Terpios zeteki (de Laubenfels).<br />

Anthosigmella varians (Duchassaing and Michelotti).<br />

Geodia papyracea n. sp.<br />

Erylus ministrongylus n. sp.<br />

VI


YALE UNIVERSITY PEABODY MUSEUM OF NATURAL HISTORY BULLETIN<br />

NO. <strong>20</strong>, 104 p., 8 PLS., 15 TEXT-FIGS., 1965.<br />

A SYSTEMATIC STUDY OF THE DEMOSPONGIAE<br />

OF PORT ROYAL, JAMAICA<br />

By GEORGE JOHN HECHTEL<br />

ABSTRACT<br />

A systematic study <strong>of</strong> the shallow water sponges <strong>of</strong> Port Royal has added 54 species to the<br />

Jamaican faunal list, <strong>of</strong> which 16 are new. One new genus, Ne<strong>of</strong>ibularia, is proposed for the<br />

preoccupied Fibularia Carter. The new species are Darwinella rosacea, Haliclona hogarthi, Adocia<br />

implexiformis, Adocia albifragilis, Pellina coeliformis, Sigmadocia caerulea, Callyspongia pallida,<br />

Halichondria magniconulosa, Microciona microchela, Microciona rarispinosa, Thalyseurypon<br />

conulosa, Ulosa hispida, Diplastrella megastellata, Geodia (Cydonium) papyracea, Erylus ministrongylus,<br />

and Corticium tetralophum.<br />

The use <strong>of</strong> field characteristics on the species level is discussed. The classification <strong>of</strong> de<br />

Laubenfels (1936a) is utilized, with several modifications. The Adociidae are transferred to the<br />

Haplosclerida, and Lundbeck's family Mycalidae is used in place <strong>of</strong> de Laubenfels' Amphilectidae.<br />

A review <strong>of</strong> the taxonomy and distribution <strong>of</strong> each species is presented. The zoogeographical<br />

affinities <strong>of</strong> the Jamaican sponge fauna are noted.<br />

1


A SYSTEMATIC STUDY OF THE DEMOSPONGIAE<br />

OF PORT ROYAL, JAMAICA*<br />

GEORGE JOHN HECHTEL<br />

INTRODUCTION<br />

The West Indian sponge fauna has not been intensively studied. Detailed<br />

regional surveys were not begun until the twentieth century, when Wilson (1902a)<br />

described a collection <strong>of</strong> Puerto Rican sponges. Arndt (1927) has studied the<br />

fauna <strong>of</strong> Curacao. In a series <strong>of</strong> papers (1936a, 1936b, 1949, 1950a, 1953a) de<br />

Laubenfels discussed the shallow water sponges <strong>of</strong> localities on the periphery <strong>of</strong><br />

the West Indies (the Dry Tortugas, in the Florida Keys; Panama; Bimini in the<br />

Bahamas; Bermuda; and the west coast <strong>of</strong> Florida, respectively.) The fauna <strong>of</strong> the<br />

Antilles is still poorly known.<br />

Jamaica was chosen for study as one <strong>of</strong> the Greater Antilles. The <strong>University</strong><br />

College <strong>of</strong> the West Indies maintains a marine station on the Jamaican south<br />

coast. The present report is a systematic study <strong>of</strong> the Demospongiae growing in<br />

less than twenty feet <strong>of</strong> water near the station.<br />

This study was submitted in 1962 to the Faculty <strong>of</strong> the Graduate School <strong>of</strong><br />

<strong>Yale</strong> <strong>University</strong> as a dissertation for the degree <strong>of</strong> Doctor <strong>of</strong> Philosophy.<br />

ACKNOWLEDGMENTS<br />

I wish to express my sincere appreciation to Dr. Willard D. Hartman, who<br />

introduced me to the study <strong>of</strong> sponges. His encouragement, advice and criticisms<br />

have been invaluable. An initial trip to Jamaica in the summer <strong>of</strong> 1959 was<br />

made possible by the Department <strong>of</strong> Zoology and the <strong>Peabody</strong> <strong>Museum</strong> <strong>of</strong> <strong>Yale</strong><br />

<strong>University</strong>. Extensive field work was made possible by an assistantship at the<br />

<strong>University</strong> College <strong>of</strong> the West Indies during the academic year 1959-60 and<br />

1960-61. Pr<strong>of</strong>essor David M. Steven <strong>of</strong> the Zoology Department at the <strong>University</strong><br />

College <strong>of</strong> the West Indies generously provided laboratory facilities at the <strong>University</strong><br />

and its marine station. Dr. Ivan Goodbody and Dr. Thomas Goreau,<br />

also <strong>of</strong> the <strong>University</strong>, contributed valuable specimens. Mr. John Howard and<br />

Dr. Patricia R. Bergquist at the <strong>Peabody</strong> <strong>Museum</strong> <strong>of</strong> <strong>Yale</strong> <strong>University</strong> gave invaluable<br />

aid in the preparation <strong>of</strong> the plates. Finally, I am deeply grateful to<br />

my wife, Florence M. Hechtel, for her aid and encouragement.<br />

METHODS<br />

Most specimens were collected by hand, after wading or diving with snorkel<br />

and face mask. Several specimens were collected in deeper water by Dr. T. F.<br />

Goreau, using an aqua-lung. Field characteristics, such as color, consistency and<br />

oscular size, were noted at the time <strong>of</strong> collection. The specimens were preserved<br />

in two changes <strong>of</strong> 75 per cent alcohol.<br />

Spicule preparations were obtained by boiling small samples in concentrated<br />

nitric acid, within a small Erlenmeyer flask. The acid was removed by a process<br />

* Published with the aid <strong>of</strong> a National Science Foundation Publication Grant, No. GN257.<br />

2


INTRODUCTION 3<br />

<strong>of</strong> washing, centrifuging, and decanting. Water and then alcohol were used in<br />

washing. The spicules were then suspended in alcohol and poured onto slides<br />

for drying and mounting.<br />

Dermal structure was studied from thin strips <strong>of</strong> the surface, removed by a<br />

razor blade or forceps. Internal morphology was examined in hand sections cut<br />

from paraffin-embedded pieces <strong>of</strong> the specimens. Sections were taken tangentially<br />

and at right angles to the surface. Longitudinal sections were made <strong>of</strong> elongate<br />

specimens. Most preparations were stained with Poinceau red or light green.<br />

Spicules are conspicuous in such sections, and spongin becomes somewhat more<br />

deeply stained than the surrounding flesh.<br />

HABITATS<br />

Port Royal is a fishing village at the tip <strong>of</strong> the Palisadoes Peninsula. The latter<br />

forms the southern shore <strong>of</strong> Kingston Harbor. The northern edge <strong>of</strong> the peninsula<br />

is fringed by mangrove swamps which begin about one mile northeast <strong>of</strong><br />

the marine station. A chain <strong>of</strong> reefs and islands (the Kingston Cays) lies <strong>of</strong>fshore<br />

from Port Royal. The geography <strong>of</strong> the area is detailed in British Admiralty<br />

maps 454 and 456. A variety <strong>of</strong> habitats could easily be reached by small boat.<br />

The principal collecting areas were:<br />

Habitat A—Port Royal Docks: Sponges were a conspicuous element <strong>of</strong> the<br />

dock and seawall community. The water was 8-10 feet deep at the base <strong>of</strong> the<br />

seawall. Many specimens were collected from the dock <strong>of</strong> the old university<br />

marine station, located just beyond the northeastern wall <strong>of</strong> Port Royal. A second<br />

collecting point was a seawall near the tide guage <strong>of</strong> the Port Royal police<br />

post on Port Royal Point.<br />

Habitat B—Port Royal Harbor: Specimens were collected in 15-<strong>20</strong> feet <strong>of</strong><br />

water, just <strong>of</strong>fshore from Port Royal, by Dr. T. F. Goreau using an aqua-lung.<br />

They were growing on coral masses.<br />

Habitat C—Rasta's wreck: A shallow, mangrove-fringed inlet opens into Port<br />

Royal harbor between the naval yard and Gallow's Point. The inlet is located at<br />

17° 56'30" N. Lat., 76° 50' W. Long. A wooden hulk, known locally as Rasta's<br />

wreck, was submerged in 4-8 feet <strong>of</strong> water at the eastern end <strong>of</strong> the inlet. Sponges<br />

covered much <strong>of</strong> its surface.<br />

Habitat D—Port Royal turtle grass bed: Turtle grass grew over much <strong>of</strong> the<br />

muddy bottom <strong>of</strong> the inlet near Rasta's wreck. The water was 3-6 feet in depth.<br />

A few species were growing on or among the turtle grass.<br />

Habitat E—Mangroves bordering inlet: The well-illuminated water was 1-2<br />

feet deep. Many mangrove roots were covered with sponges.<br />

Habitat F—Main boat channel: A rowboat channel, averaging 10 feet in<br />

width and 2-3 feet in depth, running from the northeastern corner <strong>of</strong> the inlet toward<br />

Kingston harbor. It was shaded by the tops <strong>of</strong> mangroves and had a steady<br />

current. Sponges were abundant on mangrove roots along the channel.<br />

Habitat G—Pigeon House Pile: Several species were dredged from a mussel<br />

bank near Pigeon House Pile, in Kingston harbor, at 17° 57'30" N. Lat., 76°<br />

49'30" W. Long. The sponges were growing in 10-15 feet <strong>of</strong> water.<br />

Habitat H—Shallow water at the cays: Sponges were collected under rocks<br />

and ledges in 1-6 feet <strong>of</strong> water. Study was concentrated on Drunkenman's Cay,<br />

and to a lesser extent, Maiden Cay. Drunkenman's Cay is located at 17° 54' N.<br />

Lat., 76° 30'36" W. Long. Maiden Cay is at 17° 54'30" N. Lat., 76° 48W W.<br />

Long.


4 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

Habitat I—Deeper water near the cays: Sponges were common on coral masses<br />

and gorgonian colonies in 10-15 feet <strong>of</strong> water. Most specimens were collected in<br />

an area between Lime Cay (17° 55' N. Lat., 76° 49' W. Long) and Maiden Cay.<br />

Habitat J—Offshore turtle grass beds: Several areas <strong>of</strong> sand and turtle grass<br />

were located near Port Royal. Study was concentrated on a small patch in several<br />

feet <strong>of</strong> water on the leeward side <strong>of</strong> Drunkenman's Cay.<br />

PREVIOUS JAMAICAN RECORDS<br />

Twelve published reports <strong>of</strong> Jamaican sponges, referring to ten species, have<br />

been noted in the course <strong>of</strong> the present study. The earliest record is that <strong>of</strong><br />

Higgin (1877). His Halichondria birotulata (transferred to Iotrochota by Ridley,<br />

1884) was described from Venezuela, the Bahamas, and Kingston Bay, Jamaica.<br />

De Laubenfels (1932a, p. 40) examined Jamaican specimens (those <strong>of</strong> Higgin?) in<br />

the British <strong>Museum</strong> (<strong>Natural</strong> <strong>History</strong>) collections. The species is abundant at<br />

Port Royal.<br />

Suberites angulospiculatus was described from Jamaica by Carter (1879a, p.<br />

346). Its peculiar spicules are somewhat similar in form to those <strong>of</strong> Plakortis, but<br />

are much larger. De Laubenfels (1936a, p. 162) attributed Dry Tortugas specimens<br />

to Carter's species, which he transferred to Epipolasis. De Laubenfels'<br />

sponges, with oxeas and strongyloxeas, are not even congeneric with Suberites<br />

angulospiculatus. Carter's species was not found in the present study, and its<br />

taxonomic position remains obscure.<br />

Suberites coronarius Carter (1882, p. 352) was described from Honduras, the<br />

Bahamas, and Jamaica. The species was transferred to Anthosigmella by Topsent<br />

(1918, p. 557) and placed into synonymy with A. varians by de Laubenfels (1936a,<br />

p. 13). Several specimens were collected in the present study.<br />

Spinosella maxima Dendy (1887d, p. 506; 1890, p. 365) was described from<br />

Nassau in the Bahamas. A Jamaican variety was characterized by the presence<br />

<strong>of</strong> vestigial spicules. Dendy's species, with its variety, is almost certainly conspecific<br />

with Callyspongia plicvfera, which is common near the Kingston Cays.<br />

Lendenfeld included three Jamaican records in his 1889 monograph <strong>of</strong> the<br />

Keratosa. The specimen attributed to Hircinia variabilis Schmidt was probably<br />

an example <strong>of</strong> Ircinia fasciculata. His record <strong>of</strong> Aplysina flagelliformis Carter<br />

(1886) probably refers to Verongia longissima. Both species occur at Port Royal.<br />

His Aplysina spengelii, reported from Jamaica and Ceylon, is probably a synonym<br />

<strong>of</strong> Verongia lacunosa, as suggested by Topsent (1932). It was not found at Port<br />

Royal.<br />

Spirastrella andrewsii George and Wilson (1919, p. 135) was described from<br />

North Carolina and Jamaica. De Laubenfels (1932b, p. 50) has shown that the<br />

species is conspecific with Spheciospongia vesparia. A specimen was collected in<br />

the present survey.<br />

Uliczka (1929, p. 38) described Cinachyra rhizophyta from Kingston, Jamaica.<br />

No further specimens have been found.<br />

Geodia media Bowerbank was reported by Uliczka (1929, p. 38) from Barbados,<br />

Tortugas, and Kingston. G. media is a probable synonym <strong>of</strong> the common<br />

Geodia gibberosa.<br />

CLASSIFICATION<br />

The classification <strong>of</strong> de Laubenfels (1936a) has been adopted in large part.<br />

It requires considerable revision but remains the most complete and widely-used<br />

system. Several alterations do seem necessary, however. The keratose sponges


SYSTEMATICS 5<br />

are divided into the suborders Dictyoceratina and Dendroceratina <strong>of</strong> Minchin<br />

(1900). The Adociidae are transferred to the order Haplosclerida, as discussed in<br />

the systematic section. The family Mycalidae Lundbeck (1905) is used in place<br />

<strong>of</strong> the similarly defined family Amphilectidae <strong>of</strong> de Laubenfels. The order<br />

Homosclerophorida, reconstituted by Levi (1953, 1956b), has been utilized in<br />

preference to de Laubenfels* treatment.<br />

TAXONOMIC CRITERIA<br />

In de Laubenfels* classification, genera are usually based on spicule complement<br />

or fiber composition. Species are characterized by their shape, color, consistency,<br />

spicule size ranges and details <strong>of</strong> architecture.<br />

Burton (1932, p. 377) has questioned the taxonomic value <strong>of</strong> microscleres,<br />

even on a species level. In the present study only two species were found to vary<br />

in spicule complement. The first, Sigmadocia caerulea, is a new species established<br />

for blue, sigma-rich, inshore sponges. Blue, algae-infested cay sponges with<br />

extremely rare sigmas are only provisionally considered to be conspecific with it.<br />

Gelliodes areolata provides a case <strong>of</strong> variation in the presence <strong>of</strong> microscleres between<br />

specimens which are certainly conspecific. It seems probable that most<br />

sponge species have a stable spicule complement. Spicule size ranges, considered<br />

as variable by Burton, have been relatively consistent within a species in my experience.<br />

The field characteristics <strong>of</strong> color and consistency have been invaluable in the<br />

recognition <strong>of</strong> species. The color is unfortunately lost from almost all species<br />

after preservation. The consistency <strong>of</strong> preserved sponges is usually not unlike<br />

that <strong>of</strong> the living ones. The uniformity <strong>of</strong> coloration in a large majority <strong>of</strong> species<br />

is very striking. The observed specimens <strong>of</strong> common species such as Haliclona<br />

rub ens, Iotrochota birotulata and Tedania ignis did not differ appreciably in<br />

color. The intensity <strong>of</strong> coloration does vary in many species. Algae occasionally<br />

impart a greenish tinge to a sponge. Terpios zeteki does exhibit a wide range <strong>of</strong><br />

colors, however, but can be recognized by its characteristic shape and consistency.<br />

Levi (1952, p. 37) has also emphasized the taxonomic value <strong>of</strong> color in tropical<br />

sponges.<br />

Sponges have <strong>of</strong>ten been described as extremely plastic in form, (cf., for example,<br />

Bowerbank, 1866, p. 212). Burton (1932, p. 376) and Wells et al. (1960,<br />

p. <strong>20</strong>2) have noted that most species do have a characteristic shape or range <strong>of</strong><br />

shapes. The shape can, however, be described only in general terms, such as<br />

incrusting, massive and ramose. An occasional sponge may depart from the<br />

normal. A single massive example <strong>of</strong> the usually ramose Iotrochota birotulata<br />

and a single semi-ramose example <strong>of</strong> the incrusting Microciona microchela were<br />

found.<br />

SYSTEMATICA<br />

A nomenclatorial summary is included with the description <strong>of</strong> each previously<br />

described species. The localities <strong>of</strong> collection are included within the summaries.<br />

The zoogeographical range <strong>of</strong> each species is presented under the heading <strong>of</strong><br />

distribution, using the terminology <strong>of</strong> Ekman (1953). The term spicule category<br />

refers to spicules having the same morphological form, such as sigmas. In spicule<br />

analyses, the number <strong>of</strong> measured spicules is indicated in parentheses after the<br />

range. The spicule ranges <strong>of</strong> holotypes are starred (*).<br />

Fifty-four species are discussed in the present report. One species cannot be<br />

completely classified at this time. Of the remaining 53, 16 are new. One new


6 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

genus is established. The known Jamaican sponge fauna now consists <strong>of</strong> 57<br />

species, including three (Verongia lacunosa, Suberites angulospiculatus, and<br />

Cinachyra rhizophyta) not found in the present study. The collection is deposited<br />

in the <strong>Peabody</strong> <strong>Museum</strong> <strong>of</strong> <strong>Natural</strong> <strong>History</strong>, <strong>Yale</strong> <strong>University</strong> (YPM), New<br />

Haven, Connecticut. In the text, the United States National <strong>Museum</strong> is abbreviated<br />

as USNM.<br />

The fauna <strong>of</strong> Port Royal is summarized in the following outline:<br />

CLASS DEMOSPONGIAE<br />

ORDER KERATOSA<br />

SUBORDER Dictyoceratina<br />

FAMILY SPONGIIDAE<br />

Ircinia fasciculata (Pallas), p. 8<br />

Ircinia strobilina (Lamarck), p. 10<br />

Oligoceras hemorrhages de Laubenfels, p. 11<br />

Verongia fistularis (Pallas), p. 12<br />

Verongia longissima (Carter), p. 13<br />

FAMILY DYSIDEIDAE<br />

Dysidea fragilis (Montagu), p. 14<br />

lanthella ardis de Laubenfels, p. 16<br />

SUBORDER Dendroceratina<br />

FAMILY APLYSILLIDAE<br />

Darwinella rosacea n. sp., p. 17<br />

ORDER HAPLOSCLERIDA<br />

FAMILY HALICLONIDAE<br />

Haliclona doria de Laubenfels, p. 18<br />

Haliclona rub ens (Pallas), p. 18<br />

Haliclona erina de Laubenfels, p. 19<br />

Haliclona hogarthi n. sp., p. <strong>20</strong><br />

FAMILY DESMACIDONIDAE<br />

Desmapsamma anchorata (Carter), p. 21<br />

Ne<strong>of</strong>ibularia n. gen., p. 22<br />

Ne<strong>of</strong>ibularia massa (Carter), p. 23<br />

Iotrochota birotulata (Higgin), p. 24<br />

Gelliodes areolata (Wilson), p. 25<br />

FAMILY ADOCIIDAE<br />

Adocia carbonaria (Lamarck), p. 26<br />

Adocia implexiformis n. sp., p. 27<br />

Adocia albifragilis n. sp., p. 28<br />

Pellina coeliformis n. sp., p. 29<br />

Sigmadocia caerulea n. sp., p. 30<br />

FAMILY CALLYSPONGIIDAE<br />

Callyspongia fallax Duchassaing and Michelotti, p. 31<br />

Callyspongia vaginalis (Lamarck), p. 32<br />

Callyspongia plicifera (Lamarck), p. 34<br />

Callyspongia pallida n. sp., p. 36


ORDER POECILOSCLERIDA<br />

FAMILY TEDANIIDAE<br />

Tedania ignis (Duchassaing and Michelotti), p. 37<br />

Lissodendoryx isodictyalis (Carter), p. 38<br />

Acanthacarnus souriei LeVi, p. 40<br />

FAMILY MICROCIONIDAE<br />

Microciona microchela n. sp., p. 41<br />

Microciona rarispinosa n. sp., p. 42<br />

Thalyseurypon conulosa n. sp., p. 44<br />

FAMILY MYCALIDAE<br />

Mycale laevis (Carter), p. 46<br />

Mycale microsigmatosa Arndt, p. 47<br />

Zygomycale parishii (Bowerbank), p. 48<br />

Ulosa hispida n. sp., p. 51<br />

ORDER HALICHONDRIDA<br />

FAMILY HALICHONDRIIDAE<br />

Halichondria melanadocia de Laubenfels, p. 52<br />

Halichondria magniconulosa n. sp., p. 53<br />

ORDER HADROMERIDA<br />

SYSTEMATICS 7<br />

FAMILY SPIRASTRELLIDAE<br />

Spirastrella coccinea (Duchassaing and Michelotti), p. 54<br />

Anthosigmella varians (Duchassaing and Michelotti), p. 55<br />

Spheciospongia vesparia (Lamarck), p. 57<br />

Diplastrella megastellata n. sp., p. 58<br />

FAMILY SUBERITIDAE<br />

Terpios zeteki de Laubenfels, p. 59<br />

FAMILY CLIONIDAE<br />

Cliona vermifera Hancock, p. 60<br />

Cliona viridis (Schmidt), p. 61<br />

FAMILY PLACOSPONGIIDAE<br />

Placospongia carinata (Bowerbank), p. 62<br />

ORDER EPIPOLASIDA<br />

FAMILY TETHYIDAE<br />

Tethya seychellensis (Wright), p. 65<br />

Tethya actinia de Laubenfels, p. 66<br />

Tethya sp., cf. maza Selenka, p. 67<br />

ORDER CHORISTIDA<br />

FAMILY GEODIIDAE<br />

Geodia (Geodia) gibberosa (Lamarck), p. 68<br />

Geodia (Cydonium) papyracea n. sp., p. 71<br />

Erylus ministrongylus n. sp., p. 72<br />

FAMILY CHONDRILLIDAE<br />

Chondrilla nucula Schmidt, p. 74


8 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

ORDER HOMOSCLEROPHORIDA<br />

FAMILY PLAKINIDAE<br />

Plakortis zyggompha (de Laubenfels), p. 76<br />

Corticium tetralophum n. sp., p. 77<br />

CLASS DEMOSPONGIAE Sollas, 1888<br />

ORDER KERATOSA Grant, 1861<br />

SUBORDER Dictyoceratina Minchin, 1900<br />

FAMILY SPONGIIDAE Gray, 1867,<br />

sensu de Laubenfels, 1948<br />

GENUS IRC INI A Nardo<br />

Ircinia fasciculata (Pallas) de Laubenfels<br />

Spongia fasciculata Pallas, 1766, p. 381 [neotype: Dry Tortugas, Florida (not from type<br />

locality); USNM no. 22503]; Lamarck, 1813, p. 372.<br />

[non] Spongia fasciculata Esper, 1794, p. 253 (fide Topsent, 1930.)<br />

Hircinia fasciculata, Schmidt, 1862, p. 34; Lendenfeld, 1889, p. 587; Topsent, 19<strong>20</strong>b, p.<br />

3<strong>20</strong>; idem, 1930, p. 16.<br />

Ircinia fasciculata, de Laubenfels, 1948, p. 66; idem, 1949, p. 5; idem, 1950a, p. 9; idem,<br />

1953a, p. 514; idem, 1956, p. 2; Hartman, 1955, p. 165; Wells et al., 1960, p. <strong>20</strong>6;<br />

Little, 1963, p. 34.<br />

[non] Hircinia fasciculata, Row, 1911, p. 373.<br />

Hircinia variabilis, Lendenfeld (not Schmidt, 1862), 1889, p. 557 [partim]; Wilson, 1902a,<br />

p. 405; de Laubenfels, 1936a, p. 19; idem, 1936b, p. 457; idem, 1947, p. 35.<br />

[non] Ircinia variabilis, de Laubenfels, 1950a, p. 14.<br />

Hircinia ect<strong>of</strong>ibrosa George and Wilson, 1919, p. 166 (fide de Laubenfels, 1948.)<br />

HABITAT. Common on pilings at Port Royal and on rocks near the cays.<br />

SHAPE. Lobate or branching. Many specimens consist <strong>of</strong> a cluster <strong>of</strong> laterally fused<br />

cylindrical lobes. The sponges are <strong>of</strong>ten fist-sized, but may reach 15 cm in height. Branches<br />

and projections are typically several cm in diameter. Their distal ends are somewhat<br />

rounded in outline.<br />

COLOR. In life, the upper surfaces may be yellowish-brown, reddish-brown, or reddishpurple.<br />

The interior and basal parts <strong>of</strong> the sponge are drab to cream. The oscules are<br />

dark brown to black. Preserved specimens retain some pigment, particularly around the<br />

oscular rims. The upper surfaces become olive green to gray in alcohol.<br />

CONSISTENCY. Tough, slightly compressible, difficult to cut or tear.<br />

ODOR. Fetid.<br />

SURFACE. Finely conulose. The conules on the side and upper surfaces are 0.5 to 3,<br />

<strong>of</strong>ten 1 to 2 mm, in height, and 1 to 2, or occasionally 3 mm, apart. They are less<br />

numerous near the base <strong>of</strong> the sponge. Many conules are connected to adjacent ones<br />

by ridges. A network <strong>of</strong> lines, visible to the unaided eye, radiates outward from the<br />

conules.<br />

The dark-rimmed oscules are scattered apically, sometimes in ill-defined groups. In life,<br />

they are several millimeters to nearly a centimeter in diameter. Most <strong>of</strong> them are closed<br />

in preserved specimens, with their positions indicated by darkened areas. The oscules <strong>of</strong><br />

one branching specimen are located terminally on thin processes which are about 5-6 mm<br />

in height and a few millimeters in diameter.<br />

ECTOSOME. The ectosome is a tough skin difficult to detach from the underlying flesh.<br />

A few openings, perhaps ostia, 80-100 /x in diameter, were found in surface strips. The<br />

surface contains foreign material, knobbed filaments, and spotted, knobbed filaments.<br />

Some <strong>of</strong> the debris is condensed into a network <strong>of</strong> loose tracts. The tracts are 30-150 fi


SYSTEMATICS 9<br />

in diameter; the meshes about 150 p. in width. Filaments and spotted filaments may be<br />

grouped into dense bands. The branching specimen with projecting oscules has a very<br />

course and irregular dermal network <strong>of</strong> spongin fibers which are <strong>20</strong>-60 p in diameter.<br />

George and Wilson (1919) have reported dermal fibers in their North Carolina specimens.<br />

ENDOSOME. The skeleton is a coarse fibroreticulation. The primary fibers are <strong>20</strong>-75 JM<br />

in diameter and contain an abundance <strong>of</strong> foreign material. No fibrillar pith is present.<br />

The primary fibers are grouped into loose or condensed fascicles. The fascicles ascend to<br />

the conules at intervals <strong>of</strong> 0.5-1 mm. The connective fibers are similar in size to those<br />

<strong>of</strong> the fascicles. They contain less debris than the primaries, and indeed are <strong>of</strong>ten composed<br />

entirely <strong>of</strong> spongin. Often simple in mid-length, they divide into a root-like network<br />

<strong>of</strong> narrow fibers near the fascicles. The roots may join a fascicle or run parallel to<br />

it. In many cases, the debris-filled primaries form an axial tract within a column <strong>of</strong> clear<br />

secondary fibers. Such compound fascicles have been noted in Hircinia ect<strong>of</strong>ibrosa George<br />

and Wilson, here considered as a synonym <strong>of</strong> Ircinia fasciculata. Adjacent connectives<br />

may be nearly a millimeter apart.<br />

The flesh contains some scattered debris and an abundance <strong>of</strong> filaments. The latter<br />

have a strand diameter <strong>of</strong> 3-5 p. They terminate in spherical knobs, 7-10 fi in diameter.<br />

Many strands have a similar form but a slightly larger diameter. They also differ from<br />

typical Ircinia filaments through the presence <strong>of</strong> small refractile bodies on the strands<br />

and terminal swellings. George and Wilson (1919, p. 167) noted the presence <strong>of</strong> such<br />

"spotted fibers" in Hircinia ect<strong>of</strong>ibrosa. Many strands are only partially spotted, thus<br />

providing a clear link between the spotted and typical filaments. Spotted filaments are<br />

also present in Jamaican specimens <strong>of</strong> Ircinia strobilina. De Laubenfels (1954a, p. 22)<br />

has suggested that the spotted filaments are algae which become digested and modified<br />

by the sponge.<br />

DISTRIBUTION. Tropical Atlantic America: North Carolina—George and Wilson, 1919,<br />

p. 166 (as Hircinia ect<strong>of</strong>ibrosa); de Laubenfels, 1947, p. 35 (as Hircinia variabilis)', Wells<br />

et al., 1960, p. <strong>20</strong>6. Florida—Hartman, 1955, p. 165. Dry Tortugas, Florida—de Laubenfels,<br />

1936a, p. 19 (as Hircinia variabilis). West coast, Florida—de Laubenfels, 1953a, p.<br />

514; Little, 1963, p. 34. Atlantic coast: Panama—de Laubenfels, 1936b, p. 457 (as Hircinia<br />

variabilis). Brazil—de Laubenfels, 1956, p. 2. Bermuda—de Laubenfels, 1950a, p. 9;<br />

Hartman, 1955, p. 165. Bahamas—de Laubenfels, 1949, p. 5; Hartman, 1955, p. 165.<br />

Jamaica—Lendenfeld, 1889, p. 557, partim (as Hircinia variabilis). Puerto Rico—Wilson,<br />

1902a, p. 405 (as Hircinia variabilis).<br />

Mediterranean—-Pallas, 1766, p. 381 (as Spongia fasciculata); Lamarck, 1813, p. 372<br />

(as Spongia fasciculata) (?); Adriatic—Schmidt, 1862, p. 34 (as Hircinia fasciculata) (?).<br />

DISCUSSION. The branched specimen with oscular projections and surface spongin<br />

fibers is provisionally considered to be conspecific with the other specimens. Its large,<br />

dark-rimmed oscules are <strong>of</strong> the usual Ircinia fasciculata type. The specimen resembles<br />

others in conule pattern, surface reticulation and internal architecture.<br />

The Jamaican specimens are considered to be conspecific with the fine-grained West<br />

Indian sponges attributed to Ircinia fasciculata by de Laubenfels. A few Jamaican specimens<br />

have the elongate, blunt-tipped form <strong>of</strong> /. ramosa (Keller). They differ from<br />

Keller's species in having large, dark-rimmed, fasciculata-type oscules. In addition, bundles<br />

<strong>of</strong> thick filaments and tracts <strong>of</strong> foreign material are present in the ectosome. Hartman<br />

(1955) found such a pattern in specimens <strong>of</strong> /. fasciculata but not in /. ramosa.<br />

Ircinia fasciculata was originally described by Pallas for Mediterranean specimens<br />

which have apparently been lost. The Mediterranean specimens <strong>of</strong> Esper and Lamarck<br />

have been restudied by Topsent (19<strong>20</strong>b, 1930). Topsent (1930) concluded that the two<br />

sponges were not conspecific. Lamarck's specimens, with thick filaments and debriscontaining<br />

fibers, is certainly similar in skeletal structure to the West Indian species.<br />

Schmidt (1862) used the name fasciculata for an unrecognizable, macerated, Adriatic<br />

sponge. Row's Red Sea sponge, with widely scattered conules, is certainly not conspecific<br />

with the West Indian species. The relationship <strong>of</strong> the common fine-grained West Indian<br />

Ircinia to similar sponges in other parts <strong>of</strong> the world remains obscure. Many specimens<br />

<strong>of</strong> Ircinia with low conules have been recorded as Ircinia variabilis (Schmidt), which was


10 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

considered to be a circumtropical species. Lendenfeld's Jamaican specimen is very probably<br />

an Ircinia fasciculata. De Laubenfels (1948, 1950a) concluded that most records <strong>of</strong><br />

/. variabilis Schmidt were not conspecific with the common West Indian fine-grained<br />

Ircinia. He ignored the extant Mediterranean specimens <strong>of</strong> Esper and Lamarck and<br />

unwisely chose an Ircinia from the Dry Tortugas, Florida, as the neotype <strong>of</strong> /. fasciculata<br />

(Pallas). He also listed an extensive synonymy for the species. It is by no means certain<br />

that the West Indian species under consideration has a wider distribution. De Laubenfels'<br />

synonymy certainly requires verification in every case. Vacelet (1959) has already conserved<br />

/. oros and /. dendroides Schmidt as distinct species.<br />

Levi (1952, 1959, 1960), Vacelet (1959, 1961) Sara and Siribelli (1960) and Sara (1958b,<br />

1960a, 1961a, 1961b) have used /. fasciculata for Mediterranean and West African sponges<br />

which formerly would have been called /. variabilis. The specimens <strong>of</strong> Vacelet (1959) and<br />

Sara (1961a) may be conspecific with the West Indian species. LeVi's Gulf <strong>of</strong> Guinea<br />

sponge, however, differs from the West Indian in conule pattern and color.<br />

Ircinia strobilina (Lamarck) de Laubenfels<br />

Spongia strobilina Lamarck, 1816, p. 383 [type: locality uncertain; Mus. nat. Hist. nat.<br />

Paris]; idem, 1836, p. 573.<br />

Hircinia strobilina, de Laubenfels, 1936a, p. 18.<br />

Ircinia strobilina, de Laubenfels, 1948, p. 71; idem, 1948, p. 71; idem, 1949, p. 6; idem,<br />

1950a, p. 14; idem, 1953a, p. 514; Sara, 1958a, p. 240 (?); Little, 1963, p. 35.<br />

[non] Ircinia strobilina irregularis, de Laubenfels, 1954a, p. 21.<br />

Spongia linteiformis var. ft Lamarck, 1813, p. 457 (fide Topsent, 1933a.)<br />

Hircinia gigantea, Topsent (not Lendenfeld, 1889), 1933a, p. 15.<br />

HABITAT. Common on pilings at Port Royal and on corals <strong>of</strong>fshore from the cays.<br />

SHAPE. Massive, <strong>of</strong>ten lobate or cake-shaped. Many specimens attain a height <strong>of</strong> 8-10<br />

cm, and a maximum diameter <strong>of</strong> 4-6 cm. Sponges as large as 2 or 3 feet in diameter were<br />

seen in the field.<br />

COLOR. The upper and lateral surfaces are gray to black in life and when preserved<br />

in alcohol. The basal surfaces and the interior are a dull yellowish-orange. The oscular<br />

rims are always dark gray or black.<br />

CONSISTENCY. Tough, compressible.<br />

ODOR. Unpleasant, strong.<br />

SURFACE. Coarsely conulose. The thick, blunt, <strong>of</strong>ten bifid conules are mostly 3-7 mm<br />

high, and 3-8 mm apart. Many <strong>of</strong> them are joined by high connecting ridges. The conules<br />

become lower and less frequent near the base. Fine tracts, visible to the unaided eye,<br />

radiate outward from the conules. They are joined by fine connective tracts. The oscules<br />

are 2-3 mm, occasionally 5 mm, in diameter. They occur in groups on the apices <strong>of</strong> the<br />

lobes.<br />

ECTOSOME. A tough detachable skin contains an abundance <strong>of</strong> filaments, spicule fragments<br />

and sand. Much <strong>of</strong> the debris is condensed into spongin-free tracts which are 50-<br />

150 fi in diameter. They form the above-mentioned network, with meshes frequently<br />

150-<strong>20</strong>0 /M in width. A few dermal pores, 70-150 p in diameter, were noted in the interstices.<br />

ENDOSOME. Numerous macroscopic canals traverse the interior. The skeleton consists<br />

<strong>of</strong> ascending fascicular columns <strong>of</strong> spongin fibers. The individual fibers, 30-250 /x in<br />

diameter, are heavily charged with foreign material. Short intervals <strong>of</strong> debris-free spongin<br />

occur sporadically in the skeleton. In one specimen considerable lengths <strong>of</strong> fiber have<br />

little or no foreign matter. The spongin in such intervals <strong>of</strong>ten appears laminated. The<br />

fibers never have the fibrillar pith characteristic <strong>of</strong> the subgenus Sarcotragus as redefined<br />

by Vacelet (1959). Adjacent fascicles may be more than a millimeter apart. Connective<br />

fibers are rarely present. The flesh contains scattered debris and, occasionally, large<br />

clumps <strong>of</strong> foreign material. It is permeated by a thick feltwork <strong>of</strong> Ircinia filaments which<br />

may even form thick bands. The filament strands are 3-5 /x, occasionally 7 /x, in diameter.<br />

The terminal knobs are 7-10 /x in diameter. In addition to typical filaments, the skin


SYSTEMATICS 11<br />

and interior contain slightly thicker knobbed filaments over which small refractile bodies<br />

are sparsely to thickly scattered.<br />

DISTRIBUTION. Tropical Atlantic America: Dry Tortugas, Florida—de Laubenfels,<br />

1936a, p. 18 (as Hircinia strobilina). West coast, Florida—de Laubenfels, 1953a, p. 514;<br />

Little, 1963, p. 35. Bermuda—de Laubenfels, 1950a, p. 14. Bahamas—de Laubenfels,<br />

1949, p. 6. Mediterranean: Ligurean Sea—Sara, 1958a, p. 240 (?).<br />

DISCUSSION. The Jamaican specimens are similar in appearance to Ircinia strobilina<br />

as described by de Laubenfels and photographed by Topsent. Foreign material has been<br />

recorded for fibers <strong>of</strong> I. strobilina by de Laubenfels (1950a). Topsent did not mention<br />

foreign matter in his redescription <strong>of</strong> the type. He considered Lamarck's sponge to be<br />

conspecific with Hircinia gigantea (Lendenfeld), which lacks foreign material. The Jamaican<br />

specimens are certainly conspecific with those <strong>of</strong> de Laubenfels who also studied the<br />

type. It should be noted that the type locality is unknown. Lamarck listed it as doubtfully<br />

Mediterranean. Sara (1958a) has reported the species from the Mediterranean, but he<br />

may be referring to Ircinia muscarum (Schmidt). (Cf. Vacelet 1959).<br />

De Laubenfels (1948) considered a number <strong>of</strong> Indo-Pacific species to be conspecific<br />

with 7. strobilina. New field studies and examination <strong>of</strong> the types will be needed to<br />

establish the synonymies. The Palau Islands specimen recorded as Ircinia strobilina, subspecies<br />

irregularis (Polejaeff) by de Laubenfels (1954a) differs from typical West Indian<br />

specimens by its brick red endosome and small conules.<br />

Genus OLIGOCERAS Schulze<br />

Oligoceras hemorrhages de Laubenfels<br />

(Plate I, fig. 5)<br />

Oligoceras hemorrhages de Laubenfels, 1936a, p. 16 [type: Dry Tortugas, Florida;<br />

USNM No. 22484]; idem, 1949, p. 6; idem, 1953b, p. 16; Hartman, 1955, p. 162.<br />

Oligoceras collectrix hemorrhages, de Laubenfels, 1948, p. 57.<br />

HABITAT. Common in the turtle grass area near Drunkenman's Cay. One specimen<br />

was dredged from the Pigeon House mussel bank.<br />

SHAPE. Small, lobate, spreading masses, usually less than two cm in height.<br />

COLOR. The exterior is basically brown, with red, green or purple tinges in some<br />

specimens. The interior is drab. In alcohol, the exterior becomes brown, drab, or olive<br />

green. The sponge emits a vivid red exudate when handled some time after collection.<br />

De Laubenfels has stated that death must occur before the sponge emits the pigment,<br />

(1949, 1953b). The red pigment quickly leaches out in alcohol.<br />

CONSISTENCY. Spongy, compressible.<br />

SURFACE. The surface is covered with fine conules which are less than a millimeter<br />

in height, and only 0.5-3 mm apart. The scattered oscules are 2-4 mm in diameter.<br />

ECTOSOME. Little dermal specialization. The thin skin contains a small to considerable<br />

quantity <strong>of</strong> debris. The ostia are 30-40 ^ in diameter. Both the ectosome and the endosome<br />

are heavily penetrated by blue-green filamentous algae, 5 p in diameter.<br />

ENDOSOME. The skeleton is a very irregular network <strong>of</strong> spongin-cemented masses <strong>of</strong><br />

debris. A specimen with much spicular debris has occasional debris-cored spongin fibers<br />

30-70 fj, in diameter. Such fibers are rarely found in sand-rich specimens. The latter<br />

contain clumps <strong>of</strong> coarse sand which may be as much as 700 JU, in diameter. Ascending<br />

skeletal columns typically terminate in the conules. Connecting fibers, where evident,<br />

contain foreign material. The flagellated chambers are 30-40 ju, in diameter.<br />

DISTRIBUTION. Tropical Atlantic America: Dry Tortugas, Florida—de Laubenfels,<br />

1936a, p. 16. Yucatan—Hartman, 1955, p. 162. Bahamas—de Laubenfels, 1949, p. 6.<br />

DISCUSSION. De Laubenfels (1948) somewhat doubtfully reduced Oligoceras hemorrhages<br />

to a subspecies <strong>of</strong> Oligoceras collectrix Schulze (1879). Schulze's Adriatic sponge,<br />

however, was black and had sparsely distributed conules. No exudate was reported. The<br />

skeleton had an antler-like (hirschgeweihahnlich) mode <strong>of</strong> branching. The differences<br />

warrant the retention <strong>of</strong> the West Indian sponge as a distinct species.<br />

Wilson's (1902a) Cacospongia spongeliformis <strong>of</strong> Puerto Rico was transferred to Oligo-


12 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

ceras by de Laubenfels (1948). It differs from O. hemorrhages in having an elongate,<br />

cylindrical shape. It also has less debris, particularly in the secondary fibers.<br />

Genus VERONGIA Bowerbank<br />

Verongia fistularis (Pallas) Bowerbank<br />

(Plate I, fig. 1)<br />

Spongia fistularis Pallas, 1766, p. 385 [type: American seas; repository unknown]; Esper,<br />

1794, p. 228; Lamarck, 1813, p. 435.<br />

Fistularia fistularis, Bowerbank, 1841c, p. 32; idem, 1844a, p. 37. (Genus preoccupied,<br />

fide Bowerbank, 1845.)<br />

Verongia fistularis, Bowerbank, 1845, p. 403; idem, 1864, p. <strong>20</strong>9; Hyatt, 1875, p. 402;<br />

Polejaeff, 1884a, p. 70; de Laubenfels, 1936a, p. 21; idem, 1948, p. 82; idem, 1949, p. 6;<br />

idem, 1950a, p. 17; idem, 1953a, p. 515; idem, 1956, p. 2.<br />

Luff aria fistularis, Duchassaing and Michelotti, 1864, p. 60.<br />

Aplysina fistularis, Lendenfeld, 1889, p. 422; Verrill, 1907, p. 332; Topsent, 1932, p. 71, 72.<br />

Spongia tubaeformis Lamarck, 1813, p. 435 (fide Topsent, 1932.)<br />

Verongia tenuissima Hyatt, 1875, p. 403 (fide de Laubenfels, 1948); Polejaeff, 1884a, p. 71.<br />

Verongia hirsuta var. fistularoides Hyatt, 1875, p. 403 (fide de Laubenfels, 1948.)<br />

Aplysina hirsuta, Wilson, 1902a, p. 409.<br />

HABITAT. Most <strong>of</strong> the observed specimens were growing on coral heads in 10-15 feet<br />

<strong>of</strong> water near the cays. A few specimens were found on pilings at Port Royal.<br />

SHAPE. Thick-walled, nearly cylindrical tubes <strong>of</strong> rather uniform diameter. Colonies <strong>of</strong><br />

several basally-united tubes occur. Many specimens are 10-30 cm in height. The walls<br />

may be considerably more than a centimeter in thickness, particularly near the base <strong>of</strong><br />

the sponge.<br />

COLOR. The living sponge is a bright yellow and sometimes has mustard or greenish<br />

tinges. When removed from the water the sponge soon becomes reddish and then progressively<br />

darker in color. It eventually becomes a very dark brown or black.<br />

CONSISTENCY. Toughly spongy in life, becoming stiff after death.<br />

SURFACE. The outer surface is finely conulose. The conules are usually under a millimeter<br />

in height and several millimeters apart. Many <strong>of</strong> the conules are connected to<br />

adjacent ones by low ridges. The resultant interconular depressions are shallow and<br />

inconspicuous in life and when preserved in alcohol. Dried specimens come to have a<br />

honeycombed surface with prominent saucer-shaped depressions.<br />

The wide-mouthed cloacal openings are typically several centimeters in diameter.<br />

Immediately inside the rim is a very narrow, membranous ridge. The lining <strong>of</strong> the cloacal<br />

cavity ranges from smooth to wrinkled, or even finely conulose. The cavity narrows<br />

toward the base <strong>of</strong> the sponge. A few small openings, 1 mm in diameter or less, occur<br />

on the cloacal and also the outer surface.<br />

ECTOSOME. A densely pigmented skin covers both the outer and cloacal surfaces <strong>of</strong> the<br />

sponge. No microscopic openings could be found in strips <strong>of</strong> the cloacal surface. The<br />

outer surface is pierced by numerous, scattered dermal pores. Typical ostial sizes are<br />

72 X 48, 87 X 58 and 101 x 72 /x.<br />

ENDOSOME. The skeleton is a coarse, irregular network <strong>of</strong> fibers, 70-175 JX in diameter.<br />

The fibers are amber, pithed and stratified. The pith typically occupies slightly more<br />

than I/3 <strong>of</strong> the fiber. The pith can undergo considerable variation in diameter along a<br />

fiber. Parallel fibers are 100 ^-1 mm apart.<br />

DISTRIBUTION. Tropical Atlantic America—America: Lamarck, 1813, p. 435 (as Spongia<br />

fistularis); 1813, p. 435 (as Spongia tubaeformis). Florida—Hyatt, 1875, p. 402; 1875, p.<br />

403 (as Verongia tenuissima). West coast, Florida—de Laubenfels, 1953a, p. 515. South<br />

America—Esper, 1794, p. 228 (as Spongia fistularis). Brazil—Polejaeff, 1884a, p. 71 (as<br />

Verongia tenuissima) (?); de Laubenfels, 1956, p. 2. Bermuda—Bowerbank, 1845, p. 403;<br />

Hyatt, 1875, p. 402; Verrill, 1907, p. 332 (as Aplysina fistularis); de Laubenfels, 1950a,<br />

p. 17. Bahamas—Hyatt, 1875, p. 402; de Laubenfels, 1949, p. 6. Cuba—Hyatt, 1875, p.<br />

403 (as Verongia hirsuta fistularoides). Puerto Rico—Wilson, 1902a, p. 409 (as Aplysina<br />

hirsuta). Guadeloupe—Duchassaing and Michelotti, 1864, p. 60 (as Luff aria fistularis).


SYSTEMATICS 13<br />

DISCUSSION. Verongia fistularis is a West Indian tubular species which undergoes a<br />

marked color change with death. The large dimensions <strong>of</strong> the tubes suggest that the<br />

slender, solid West Indian specimens <strong>of</strong> Verongia are not conspecific with fistularis.<br />

De Laubenfels (1948) has placed Luffaria sebae Duchassaing and Michelotti (1864,<br />

p. 59), and Aplysina praetexta Hyatt (1875, p. 405) into synonymy with Verongia fistularis.<br />

The first, to judge from the original description, is more likely to be V. lacunosa.<br />

The second was very inadequately described and apparently had a cup-like shape. Hyatt's<br />

Verongia hirsuta included both solid and tubular sponges with a rough, probably macerated<br />

surface. Aplysina fenestrata, also listed as a synonym <strong>of</strong> V. fistularis by de Laubenfels,<br />

is clearly congeneric with it, but not conspecific. As described by both Carter (1882)<br />

and Wilson (1902a), the internal skeleton has a peculiar honeycombed architecture.<br />

Verongia longissima (Carter) de Laubenfels<br />

Aplysina longissima Carter, 1882, p. 271 [type: Nassau, Bahamas; Brit. Mus. (Nat. Hist.)]<br />

Verongia longissima, de Laubenfels, 1936a, p. 21, 23; idem, 1948, p. 85; idem, 1953a, p.<br />

515; idem, 1956, p. 2, 3; Little, 1963, p. 35.<br />

Aplysina flagelliformis, Lendenfeld (not Carter, 1886), 1889, p. 412 \partim]; Wilson,<br />

1902a, p. 406 (including var. rugosa, p. 407.)<br />

HABITAT. The first specimen was collected by Dr. T. F. Goreau <strong>of</strong> the <strong>University</strong> <strong>of</strong><br />

the West Indies in <strong>20</strong> feet <strong>of</strong> water <strong>of</strong>fshore from Port Royal. He found a second specimen<br />

in shallow water at a point eleven miles east <strong>of</strong> Kingston in St. Thomas parish.<br />

SHAPE. The first (i.e. Port Royal) sponge is a solid cylinder <strong>20</strong> cm in length and about<br />

1 cm in diameter. It arises from a basal crust which has an area <strong>of</strong> several square centimeters.<br />

The second specimen is a cluster <strong>of</strong> erect, anastomosing branches with a maximum<br />

height <strong>of</strong> 18 cm.<br />

COLOR. When received, the first specimen was reddish in color; the second had a<br />

yellowish to reddish coloration. The Port Royal sponge is grayish-brown in alcohol. The<br />

second specimen, which is dry, is now a dark reddish-brown.<br />

CONSISTENCY. Both sponges were tough and stiff when received.<br />

SURFACE. Finely conulose. The conules are seldom 1 mm and <strong>of</strong>ten only 100-300 //, in<br />

height. Adjacent conules are usually 1 mm apart and are <strong>of</strong>ten connected by low ridges.<br />

The skeleton <strong>of</strong> the dried, partially macerated St. Thomas sponge projects above the<br />

flesh over much <strong>of</strong> the surface. The oscules are 1-2 mm in diameter. They are scattered<br />

irregularly on the first and partly arranged in linear rows in the second example. Some<br />

oscules are in depression in the latter sponge. The depressions <strong>of</strong>ten have conules on<br />

their walls and probably resulted from drying.<br />

ECTOSOME. A thin outer skin is present in both examples. Dermal pores are scattered<br />

over the surface at intervals <strong>of</strong> about 75 p. Typical ostial sizes are 58 X 43, 87 X 65,<br />

116 x 58, and 145 x 58 p. Spongin fibers occasionally run parallel to and just beneath<br />

the surface in the interconular ridges.<br />

ENDOSOME. In both examples, the skeleton is a coarse-meshed reticulation <strong>of</strong> stratified,<br />

amber, pithed spongin fibers. Fiber diameter ranges between 40-115 //,. Parallel fibers are<br />

300 /A to 1 mm apart. The finely granular pith usually occupies y3 to 14 <strong>of</strong> the diameter.<br />

DISTRIBUTION. Tropical Atlantic America: Dry Tortugas, Florida—de Laubenfels,<br />

1936a, p. 21, 23. West coast, Florida—de Laubenfels, 1953a, p. 515; Little, 1963, p. 35.<br />

Brazil—de Laubenfels, 1956, p. 2, 3. Bahamas—Carter, 1882, p. 271 (as Aplysina longissima).<br />

Jamaica—Lendenfeld, 1889, p. 412, partim (?). Puerto Rico—Wilson, 1902a, p. 406<br />

(as Aplysina flagelliformis and A. flagelliformis var. rugosa).<br />

DISCUSSION. The Jamaican specimens are similar in color and consistency to Aplysina<br />

longissima Carter. The peculiar star-like surface knots <strong>of</strong> Carter's sponge are not present.<br />

They may have been the result <strong>of</strong> maceration. My specimens are almost certainly conspecific<br />

with Verongia longissima as understood by de Laubenfels (1936a, 1948). He stressed<br />

particularly the relative lack <strong>of</strong> color change in the species. The sponge is described as<br />

yellow in life, carmine soon after collection, and dull gray in alcohol. My specimen when<br />

received may have already begun to alter in color.<br />

Carter described three species, under as many generic names, which are closely related


14 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

to, if not conspecific with, V. longissima. The first, Aplysina cauliformis Carter (1882, p.<br />

270), <strong>of</strong> the Bahamas, is a solid cylindrical sponge which differs from V. longissima only<br />

in having "flaccid" spongin fibers and "subpenicillate" rather than star-like surface fiber<br />

knots. De Laubenfels (1948, p. 87) considered A. cauliformis to be a synonym <strong>of</strong> Verongia<br />

aurea. The pinkish-brown color reported by Carter is certainly inconsistent with de Laubenfels'<br />

statement (1948, p. 85) that aurea "has to an even greater extent than fistularis<br />

the property <strong>of</strong> changing color upon death." Aplysina cauliformis is more likely to be<br />

conspecific with V. longissima. It would, in that case, take precedence over the latter<br />

name. Until the types can be re-examined, it is certainly convenient to retain the name<br />

V. longissima for the sponges reported by de Laubenfels from several localities. We can<br />

provisionally characterize the Aplysina cauliformis <strong>of</strong> Carter by its flaccid skeleton.<br />

Luff aria cauliformis Carter (1882, p. 268) <strong>of</strong> Antigua and Nassau was described as a<br />

typically black, stiff, cylindrical sponge, containing stiff, fragile fibers. Carter reported<br />

that Luff aria cauliformis and Aplysina cauliformis <strong>of</strong>ten grew in "conjunction" at Nassau.<br />

The pale varieties <strong>of</strong> L. cauliformis are very similar in external appearance to A. cauliformis.<br />

The distinction between Luffaria and Aplysina as understood by Carter was based<br />

largely on the presence <strong>of</strong> a continuous pith in the former and a supposedly discontinuous<br />

pith in the latter genus (cf. Carter, 1875). We can distinguish Verongia longissima<br />

from Luffaria cauliformis on the basis <strong>of</strong> the black color reported for the dried typical<br />

variety <strong>of</strong> the latter species. A dried, black Verongia had very probably undergone considerable<br />

color change after death.<br />

Hircinia flagelliformis Carter (1886a, p. 372) <strong>of</strong> southern Australia is also similar in<br />

form to V. longissima. Carter himself considered it "just possible that the two are the<br />

same." (p. 373.) Lendenfeld, who examined the type <strong>of</strong> flagelliformis, concluded that the<br />

Australian sponge was indeed conspecific with Carter's earlier species. He retained the<br />

later-published name in violation <strong>of</strong> the rules <strong>of</strong> priority. Lendenfeld also assigned a<br />

Jamaican sponge in the collections <strong>of</strong> the British <strong>Museum</strong> to Aplysina flagelliformis<br />

(Carter). De Laubenfels (1948, p. 87) considered Hircinia flagelliformis Carter to be<br />

unrecognizable. He (p. 86) curiously included Lendenfeld's Australian record, based on<br />

Carter's sponge, in the distribution <strong>of</strong> V. longissima. Two factors weigh against the<br />

acceptance <strong>of</strong> the conspecificity <strong>of</strong> V. longissima and the Australian species. The geographical<br />

gap is wide, and the fibers may differ in structure. Carter described Hircinia<br />

flagelliformis as containing sand-cored primary and transparent lateral fibers. The<br />

Hirciniosa <strong>of</strong> his classification (1875, p. 136) were characterized by just such a distinction<br />

between the fibers. Lendenfeld neither mentioned nor denied the existence <strong>of</strong> two<br />

types <strong>of</strong> fibers.<br />

FAMILY DYSIDEIDAE Gray, sensu de Laubenfels, 1948<br />

GENUS DYSIDEA Johnston<br />

Dysidea fragilis (Montagu) Johnston<br />

Spongia fragilis Montagu, 1818, p. 114 [type: Devon, England; Brit. Mus. (Nat. Hist.) (?)].<br />

Duseideia fragilis, Johnston, 1842, p. 187 (corrected to Dysidea on p. 251.); Burton, 1932,<br />

p. 341.<br />

Dysidea fragilis, Johnston, 1842, p. 251; Bowerbank, 1864, p. 211; idem, 1866, p. 381;<br />

idem, 1874, p. 175; idem, 1882, p. 188; Hyatt, 1875, p. 545; Carter, 1876, p. 232;<br />

Burton, 1934, p. 583; idem, 1936a, p. 26; idem, 1937, p. 41; idem, 1956, p 137;<br />

idem, 1959b, p. 51; de Laubenfels, 1936a, p. 27; idem, 1948, p. 137; idem, 1950a, p.<br />

22; idem, 1950b, p. 9; idem, 1951b, p. 213; idem, 1953a, p. 515; idem, 1954a, p. 35;<br />

idem, 1955, p. 138; Rao, 1941, p. 463; Alander, 1942, p. 17; Arndt, 1943, p. 343;<br />

Sara, 1958a, p. 241; idem, 1958b, p. 274; idem, 1960a, p. 468; idem, 1960b, p. 263;<br />

Sara and Siribelli, 1960, p. 87; Sara, 1961b, p. 15; LeVi, 1959, p. 138; Vacelet, 1959, p.<br />

67; idem, 1960, p. 270; idem, 1961, p. 43; Wells et al., 1960, p. <strong>20</strong>6; Bergquist, 1961,<br />

p. 211.<br />

Spongelia fragilis, Schmidt, 1870, p. 27; Lendenfeld, 1889, p. 660 [partim]; Topsent, 1891,<br />

p. 533; idem, 1891c, p. 127; idem, 1892, p. 134; idem, 1894, p. 43; idem, 1896, p. 123;


SYSTEMATICS 15<br />

idem, 1897, p. 482; idem, 1902b, p. 329; idem, 1925b, p. 452; Topsent and Olivier,<br />

1943, p. 6; Dendy, 1905, p. <strong>20</strong>8; idem, 1916a, p. 139; Verrill, 1907, p. 332; Stephens,<br />

1912, p. 39; idem, 1917, p. 14; Hentschel, 1912, p. 447; idem, 1929, p. 994; Hernandez,<br />

1916, p. 38; idem, 1917, p. 32; Wilson, 1925, p. 476 (var. fasciculata); Burton, 1933,<br />

p. 242; Br0ndsted, 1934, p. 25.<br />

Halichondria areolata Johnston, 1842, p. 121 (fide Bowerbank, 1866.)<br />

Spongelia incrustans Schmidt, 1862, p. 29 (fide Schmidt, 1864, and Burton, 1934.)<br />

Spongelia pallescens Schmidt, 1862, p. 30 {fide Burton, 1934); idem, 1864, p. 28; idem,.<br />

1870, p. 27; Carter, 1876, p. 232; Schulze, 1878a, p. 150, 154; Polejaeff, 1884, p. 42;<br />

Wilson, 1902a, p. 410; Topsent, 1891b, p. 13; idem, 1925a, p. 715.<br />

[non] Spongelia pallescens, de Laubenfels, 1935b, p. 327 (fide de Laubenfels, 1950b.)<br />

Dysidea coriacea Bowerbank, 1874, p. 175 (fide Stephens, 1912, and Burton, 1934); idem,<br />

1882, p. 188.<br />

Spongelia elastica var. massa and var. lobata Lendenfeld, 1889, p. 658.<br />

Spongelia elastica var. lobosa Dendy, 1905, p. <strong>20</strong>8.<br />

HABITAT. Common on pilings and mangrove roots at Port Royal. Also found in <strong>of</strong>fshore<br />

turtle grass beds.<br />

SHAPE. Incrusting to massive or lobate. Many specimens are fist-sized.<br />

COLOR. Living specimens vary in color from a dull gray to a gray-blue or a pale purple.<br />

In alcohol the sponges become drab, with purplish specimens retaining tinges <strong>of</strong> that<br />

color.<br />

CONSISTENCY. S<strong>of</strong>t, compressible.<br />

SURFACE. Finely conulose. The conules are 0.5 to 1 mm in height and 1-2 mm apart.<br />

A delicate, nearly transparent skin covers the surface. It is pierced by oscules which are<br />

usually 2-3 mm in diameter. In life, the oscules are <strong>of</strong>ten rimmed by transparent collars<br />

several millimeters in height.<br />

ECTOSOME. A detachable skin contains a small amount <strong>of</strong> foreign matter. The dermal<br />

pores are <strong>20</strong>-40 jx in diameter. Groups <strong>of</strong> 6-12 ostia are separated by dense bands <strong>of</strong> flesh<br />

which are <strong>20</strong>-35 p in diameter.<br />

ENDOSOME. The skeletal fibers form a very irregular reticulation. In the interior, parallel<br />

fibers may be as much as one millimeter apart. Considerable amounts <strong>of</strong> debris,<br />

including large clumps, may be present in the flesh. Larger fibers, particularly in a sandrich<br />

specimen, are columns <strong>of</strong> debris cemented by spongin. Smaller fibers, 30-150 /*, in<br />

diameter, are composed <strong>of</strong> spongin which is lightly to heavily cored with foreign material.<br />

The quantity <strong>of</strong> sand or spicule fragments present in the sponge probably depends on<br />

the available material. The sandiest specimen was growing in a sandy turtle grass area<br />

at Drunkenman's Cay. Specimens collected from pilings and mangrove roots contain<br />

largely spicular debris. The interior contains numerous, narrow, elongate flagellated<br />

chambers. Typical sizes are 58 x 29, 65 x 50, and 87 X 72 /*,. Large embryos are present<br />

in specimens collected in September and December.<br />

DISTRIBUTION. Tropical Atlantic America: North Carolina—Wells et al., 1960, p. <strong>20</strong>6.<br />

Florida—Schmidt, 1870, p. 27 (as Spongelia pallescens); Hyatt, 1875, p. 545. Dry Tortugas,<br />

Florida—de Laubenfels, 1936a, p. 27. West coast, Florida—de Laubenfels, 1953a,<br />

p. 515. Brazil—Polejaeff, 1884, p. 42 (as Spongelia pallescens); Lendenfeld, 1889, p. 660<br />

(as Spongelia fragilis). Bermuda—Verrill, 1907, p. 332 (as Spongelia fragilis); de Laubenfels,<br />

1950a, p. 22. Antilles—Schmidt, 1870, p. 27 (as Spongelia pallescens). Puerto Rico—<br />

Wilson, 1902a, p. 410 (as Spongelia pallescens).<br />

European Boreal Atlantic—North Atlantic: Carter, 1876, p. 232; Topsent, 1892,<br />

p. 134 (as Spongelia fragilis). Iceland—Schmidt, 1870, p. 27 (as Spongelia pallescens);<br />

Carter, 1876, p. 232 (as Spongelia pallescens); Burton, 1959b, p. 51. Ireland—Stephens,<br />

1912, p. 39 (as Spongelia fragilis); 1917, p. 14 (as Spongelia fragilis). Britain—Montagu,<br />

1818, p. 114 (as Spongia fragilis); Johnston, 1842, p. 187 (as Duseideia) and p. 251<br />

(as Dysidea); Johnston, 1842, p. 121 (as Halichondria areolata); Bowerbank, 1864, p. 211;<br />

1866, p. 381; 1874, p. 175; 1882, p. 188; Bowerbank, 1874, p. 175, (as Dysidea coriacea);<br />

1882, p. 188 (as Dysidea coriacea). Sweden—Alander, 1942, p. 17. Germany—Arndt, 1943,<br />

p. 343. Rosc<strong>of</strong>f, France—Topsent, 1891, p. 533, (as Spongelia fragilis). Atlantic coast,


16 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

France—Topsent, 1891c, p. 127 (as Spongelia fragilis). Atlantic coast, Spain—Hernandez,<br />

1917, p. 32 (as Spongelia fragilis).<br />

Mediterranean—Atlantic—Mediterranean coast: Spain—Hernandez, 1916, p. 38 (as<br />

Spongelia fragilis). Mediterranean coast, France—Topsent, 1896, p. 123 (as Spongelia<br />

fragilis); 1925b, p. 17 (as Spongelia fragilis); Topsent and Olivier, 1943, p. 6 (as Spongelia<br />

fragilis); Vacelet, 1959, p. 67; 1960, p. 270. Ligurean Sea, Italy—Sara, 1958a, p. 241.<br />

Naples, Italy—Topsent, 1925a, p. 715 (as Spongelia pallescens); Sara, 1958b, p. 274;<br />

1960a, p. 468; 1960b, p. 263; 1961b, p. 15; Sara and Siribelli, 1960, p. 87. Adriatic-<br />

Schmidt, 1862, p. 29 (as Spongelia incrustans); 1862, p. 30 (as Spongelia pallescens);<br />

Schulze, 1878a, p. 150, 154 (as Spongelia pallescens). Corsica—Vacelet, 1961, p. 43.<br />

Mediterranean coast, Egypt, Burton, 1936a, p. 26. Tunisia—Topsent, 1894a, p. 43 (as<br />

Spongelia fragilis). Algeria—Topsent, 1902b, p. 328 (as Spongelia fragilis). Black Sea—<br />

de Laubenfels, 1951b, p. 213.<br />

Tropical West Africa—Topsent, 1891b, p. 13 (as Spongelia pallescens); Burton, 1956,<br />

p. 137; Levi, 1959, p. 138.<br />

South Atlantic: Ascension Island—Burton, 1932, p. 341 (as Duseideia fragilis).<br />

South Africa—Burton, 1933, p. 242 (as Spongelia fragilis). Indian Ocean—Zanzibar,<br />

East Africa—Hyatt, 1875, p. 545; Burton, 1959a, p. 272. South Arabian coast—Burton,<br />

1959a, p. 272. Mauritius—Lendenfeld, 1889, p. 660 (as Spongelia fragilis). Okhamandal,<br />

India—Dendy, 1916a, p. 139 (as Spongelia fragilis). Madras, India—Burton, 1937, p. 41.<br />

Ceylon—Dendy, 1905, p. <strong>20</strong>8 (as Spongelia fragilis); 1905, p. <strong>20</strong>8 (as Spongelia elastica<br />

var. lobosa); Rao, 1941, p. 463. Indo-Malaya—Malaya—Rao, 1941, p. 463. Straits <strong>of</strong><br />

Malacca—Lendenfeld, 1889, p. 658 (as Spongelia elastica varieties massa and lobata).<br />

East Indies—Topsent, 1897, p. 482 (as Spongelia fragilis); Hentschel, 1912, p. 447 (as<br />

Spongelia fragilis); Br0ndsted, 1934, p. 25 (as Spongelia fragilis).<br />

Australia—Lendenfeld, 1889, p. 660 (as Spongelia fragilis); 1889, p. 658 (as Spongelia<br />

elastica, varieties massa and lobata). Great Barrier Reef—Burton, 1934, p. 583. Chatham<br />

Islands—Bergquist, 1961, p. 211.<br />

West Central Pacific—de Laubenfels, 1954a, p. 35; 1955, p. 138.<br />

DISCUSSION. The circumtropical Dysidea fragilis extends into cold temperate and even<br />

sub-Arctic waters in the northeastern Atlantic. Although it seems unlikely that all <strong>of</strong><br />

the populations are conspecific, present evidence does not permit a division into species<br />

or subspecies. The extensive synonymies <strong>of</strong> Burton (1934) and de Laubenfels (1948)<br />

require verification. Bowerbank (1866) and Burton (1934) have discussed intraspecific<br />

variation in Dysidea fragilis.<br />

De Laubenfels (1936a) described Dysidea etheria from the West Indies. It differs from<br />

the Jamaican sponge in having a brilliant sky-blue color and a coarser surface. Its primary<br />

and secondary fibers enclosed different types <strong>of</strong> foreign matter.<br />

GENUS IANTHELLA Gray<br />

Ianthella ardis de Laubenfels<br />

lanthella ardis de Laubenfels, 1950a, p. 31 [type: Bermuda; Brit. Mus. (Nat. Hist.) reg.<br />

no. 1950.5.23.1]; idem, 1953a, p. 516; Little, 1963, p. 36.<br />

lanthella basta, de Laubenfels (not Pallas, 1766), 1936a, p. 31 (fide de Laubenfels, 1950a).<br />

HABITAT. A single specimen was collected by Dr. T. F. Goreau <strong>of</strong> the <strong>University</strong> <strong>of</strong><br />

the West Indies, in shallow water, eleven miles east <strong>of</strong> Kingston (in St. Thomas parish).<br />

SHAPE. Massive, attaining a maximum thickness <strong>of</strong> 5 cm.<br />

COLOR. The sponge had a bluish-purple exterior, and a yellowish interior when received<br />

on shore. In alcohol, the entire specimen is purplish-black, the exterior being<br />

darkest in color.<br />

CONSISTENCY. The sponge is stiff and exhibits little compressibility. When cut, it has<br />

the consistency <strong>of</strong> cheese.<br />

SURFACE. Smooth to the touch, but uneven. Low, blunt conules, 1-3 mm in height, are<br />

scattered over the surface. They are joined to each other by flattened ridges and raised


SYSTEMATICS 17<br />

plateaux. The resultant depressions may be areas up to 5 mm in diameter, or merely<br />

narrow furrows. The sparsely distributed oscules are 2-3 mm in diameter.<br />

ECTOSOME. The detachable skin is strongly pigmented. It is pierced by dermal pores,<br />

that vary in size from 40 x 30 to 90 X 40 /A. Groups <strong>of</strong> 5-10 ostia are located in irregularly<br />

polygonal areas, 150-2<strong>20</strong> ^ in maximum width. They are separated by broad dense<br />

bands <strong>of</strong> flesh.<br />

ENDOSOME. The interior is traversed by canals which <strong>of</strong>ten reach a centimeter in diameter.<br />

The skeleton is coarse-meshed and irregular. Adjacent fibers are usually several<br />

millimeters apart. They branch acutely and occasionally anastomose. The spongin fibers<br />

are yellow, laminated and pithed. Fiber diameter varies between 300-600 [i, with the<br />

pith accounting for 35-90 per cent <strong>of</strong> the total. The amount <strong>of</strong> pith is not constant even<br />

for fibers <strong>of</strong> similar diameter. The pith contains a granular material and also small cells,<br />

7-10 n in diameter, <strong>of</strong> unknown function. Debris, largely in the form <strong>of</strong> coarse sand<br />

grains, is sparingly distributed in the flesh, but is absent from the fibers.<br />

DISTRIBUTION. Tropical Atlantic America: Dry Tortugas, Florida—de Laubenfels,<br />

1936a, p. 31 (as Ianthella basta). West coast, Florida—de Laubenfels, 1953a, p. 516; Little,<br />

1963, p. 36. Bermuda—de Laubenfels, 1950a, p. 31.<br />

DISCUSSION. De Laubenfels has recorded the color in life as yellow, yellow-green or<br />

grass-green. He has noted a change to blue and then to purple after death. My specimen<br />

had apparently begun such a transformation. Ianthella ianthella de Laubenfels, 1949,<br />

another West Indian representative <strong>of</strong> the genus, differs from the present species by its<br />

carmine red color in life, more dendritic skeleton, peculiarly knobbed conules and much<br />

more delicate fibers.<br />

SUBORDER Dendroceratina Minchin, 1900<br />

FAMILY APLYSILLIDAE Vosmaer, sensu de Laubenfels, 1948<br />

GENUS DARWINELLA Muller<br />

Darwinella rosacea n. sp.<br />

HOLOTYPE. YPM 5032. Mangrove boat channel, July 13, 1961.<br />

HABITAT. The sponge is common on mangrove roots and shells.<br />

SHAPE. A small thin crust, seldom exceeding 0.5 mm in thickness. Most specimens are<br />

only 1-2 square cm in area.<br />

COLOR. The living sponge is pink. In alcohol, it becomes pinkish-brown or beige.<br />

CONSISTENCY. S<strong>of</strong>t.<br />

SURFACE. Conulose. The slender conules rise about 0.5 mm above the surrounding<br />

surface. They are one to several millimeters apart. No oscular openings were seen. A<br />

network <strong>of</strong> bands, visible under a magnification <strong>of</strong> <strong>20</strong> X, radiates outward from the<br />

conules.<br />

ECTOSOME. A delicate detachable dermal membrane is present. The dermal pores are<br />

<strong>20</strong>-30 fj, in diameter. They are typically located in groups <strong>of</strong> 6 to 12. The partitions<br />

between adjacent pores are 7-15 p in diameter. The previously mentioned surface bands,<br />

30 fx in diameter, enclose the pore groups.<br />

ENDOSOME. The spongin fibers run obliquely from their basal expansions up to the<br />

conules. The fibers branch infrequently and never anastomose with adjacent parts <strong>of</strong><br />

the skeleton. They are 40-100 ^ in diameter just above the base, and gradually diminish<br />

in size toward their distal end. The fibers consist <strong>of</strong> a longitudinally striated cortex and<br />

a central pith. The latter usually occupies somewhat more than one half <strong>of</strong> the total<br />

fiber diameter. At irregular intervals, the pith is crossed by transverse, distally convex<br />

lines.<br />

The spongin spicules are triactinal, with sharp-pointed rays. They lie free in the flesh,<br />

near the base <strong>of</strong> the sponge. The rays are only 130-276 ^ in length. Their basal diameter<br />

ranges between 10-17 p. The rays may be nearly straight, with slightly irregular outlines,<br />

or bent near their proximal or distal ends.<br />

Much <strong>of</strong> the interior is occupied by closely packed, sac-like flagellated chambers.<br />

Typical sizes include 87 X 43, 94 x 72, and 108 X 87 /x. Specimens collected in May,<br />

July and August contain embryos near the substratum.


18 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

DISCUSSION. TWO species <strong>of</strong> Darwinella have been reported from Tropical Atlantic<br />

America. Darwinella joyeuxi Topsent, 1889, is a cylindrical sponge with long-rayed<br />

spicules. D. millleri (M. Schultze) Miiller is a yellow sponge with multi-rayed spicules.<br />

The Darwinella millleri recorded by de Laubenfels (1950a) from Bermuda is a large<br />

ramose sponge with long-rayed triactinal spicules. It is certainly conspecific with neither<br />

the Jamaican sponge nor the D. millleri <strong>of</strong> Miiller and Schulze.<br />

The Jamaican sponge is most similar to D. intermedia Topsent (1894e) <strong>of</strong> France.<br />

Topsent's species differs in having a yellow color and spicules with markedly flexuous<br />

rays <strong>of</strong> a rather uniform small size (100-150 p). Taking the differences and wide geographical<br />

gap into consideration, the Jamaican sponge is unlikely to be conspecific with<br />

D. intermedia.<br />

ORDER HAPLOSCLERIDA Topsent 1928<br />

FAMILY HALICLONIDAE de Laubenfels 1932b<br />

GENUS HALICLONA Grant<br />

Haliclona doria de Laubenfels<br />

(Plate I, fig. 4)<br />

Haliclona doria de Laubenfels, 1936b, p. 458 [type: Fort Randolph, Panama; USNM<br />

no. 22245.]<br />

HABITAT. Several specimens were collected at Gun Cay. The species was very abundant<br />

in the shallow water turtle grass bed at the Port Royal end <strong>of</strong> the Port Royal-Kingston<br />

mangrove boat channel.<br />

SHAPE. A typical specimen is a complex <strong>of</strong> repent, anastomosing, cylindrical to ovoid<br />

branches which are 1-1.5 cm in diameter. A colony may extend over several feet <strong>of</strong> turtle<br />

grass. Gun Cay specimens are thicker, with branches <strong>of</strong>ten several centimeters in<br />

diameter.<br />

COLOR. In life, the upper surfaces are brownish-yellow to dark brown with a purplish<br />

tinge. Under-surfaces are light brown to cream. The interior is yellowish. In alcohol,<br />

the specimens are drab.<br />

CONSISTENCY. Stiff, easily broken into pieces.<br />

SURFACE. Even. The scattered oscules, 1-4 mm in diameter, <strong>of</strong>ten have low rims.<br />

ECTOSOME. No dermal specialization. The vertical bundles <strong>of</strong> the interior merely come<br />

to an end, forming a microhispid surface.<br />

ENDOSOME. The skeleton is a rather symmetrical framework <strong>of</strong> spicule tracts. The<br />

meshes are square to rectangular, with a maximum width <strong>of</strong> 70-240 p. The bundles are<br />

loosely organized, 30-60 ^ in diameter, and occasionally bound at nodes by small amounts<br />

<strong>of</strong> spongin. In addition, many spicules are scattered in the flesh.<br />

SPICULES. Oxeas, stout, straight to slightly curved, typically hastate, with the points<br />

beginning at about 1 1/2 diameters from the end, 140-188 X 3-12 /A. Individual analyses,<br />

lengths in microns, 50 spicules each: 140-176; 152-174; 140-176; 152-188; 140-176; 140-176.<br />

DISTRIBUTION. Tropical Atlantic America; Atlantic coast <strong>of</strong> Panama—de Laubenfels,<br />

1936b, p. 458.<br />

DISCUSSION. The dark brown color and pasteboard consistency <strong>of</strong> Haliclona doria is<br />

unusual for the genus. H. longleyi de Laubenfels, 1932a, <strong>of</strong> the West Indies does resemble<br />

H. doria in shape and consistency. It differs in having a yellowish-green color in life. Its<br />

spicules are more fusiform and less robust than those <strong>of</strong> H. doria. The Gun Cay specimens<br />

are identical in structure and spiculation with the inshore specimens. They have,<br />

however, a coarsely porous surface and thicker, more irregularly shaped branches. They<br />

are therefore only provisionally considered as conspecific with Haliclona doria.<br />

Haliclona rub ens (Pallas) de Laubenfels<br />

Spongia rubens Pallas, 1766, p. 389 [type: American seas; repository unknown]; Duchassaing<br />

and Michelotti, 1864, p. 41.<br />

Pachychalina rubens, Schmidt, 1870, p. 37; Wilson, 1902a, p. 392.<br />

Chalina rubens, Carter, 1882, p. 276 [partim.]


SYSTEMATICS 19<br />

Haliclona rubens, de Laubenfels, 1932a, p. 59; idem, 1936a, p. 40; idem, 1949, p. 9; idem,<br />

1953a, p. 519; Burton, 1954, p. 223; Hartman, 1955, p. 167; Little, 1963, p. 39.<br />

Amphimedon arhorescens Duchassaing and Michelotti, 1864, p. 79 {fide Wilson, 1902a.)<br />

HABITAT. The species is common on coral outcroppings near the cays.<br />

SHAPE. Some specimens are irregularly massive but most are ramose. In the latter<br />

case, the erect branches may reach a height <strong>of</strong> more than one foot and a diameter <strong>of</strong><br />

1-2 cm.<br />

COLOR. A very constant dark red which becomes somewhat darker in preserved and<br />

dried specimens.<br />

CONSISTENCY. Spongy, compressible.<br />

SURFACE. Even, punctiform. The scattered oscules are 2-5 mm in diameter.<br />

ECTOSOME. No dermal specialization.<br />

ENDOSOME. The skeleton is an irregular network <strong>of</strong> fibers and tracts, typically with<br />

rounded meshes. Numerous interstitial spicules are present. Spongin varies in amount<br />

within the skeleton <strong>of</strong> an individual. Many fibers <strong>of</strong> 30-45 /x diameter seem to lack it<br />

entirely. In adjacent parts <strong>of</strong> the skeleton spongin may form either a cementing layer<br />

or the greater part <strong>of</strong> the fiber. In the latter case, the fibers contain a multispicular core<br />

and may reach 70-90 ju, in total diameter.<br />

SPICULES. Oxeas, blunt to hastate, mostly curved, 117-163 x 3-8 p. As noted by Hartman<br />

(1955), many <strong>of</strong> the blunt spicules are actually styles or strongyles. Their frequency<br />

varies in different specimens. In one specimen, nearly <strong>20</strong> per cent <strong>of</strong> the spicules are<br />

strongyles. Some strongyles have irregular lumps in mid-shaft. Individual analyses, oxeas,<br />

lengths in microns, 50 spicules each: 117-146; 117-140; 123-163; 117-146.<br />

DISTRIBUTION. Tropical Atlantic America: American Seas—Pallas, 1776, p. 389 (as<br />

Spongia rubens); Florida—Schmidt, 1870, p. 37 (as Pachychalina rubens); Dry Tortugas,<br />

Florida—de Laubenfels 1932a, p. 59 and 1936a, p. 40; West Coast, Florida—de Laubenfels,<br />

1953a, p. 519; West Coast, Florida—Little, 1963, p. 39; Yucatan—Hartman,<br />

1955, p. 167; Bahamas—Carter, 1882, p. 276 (as Chalina rubens), de Laubenfels, 1949, p.<br />

9; Cuba—Duchassaing and Michelotti, 1864, p. 41 (as Spongia rubens); Puerto Rico—<br />

Wilson, 1902a, p. 392 (as Pachychalina rubens); Gorda Cay, Mosquito Bank, West Indies<br />

—Burton, 1954, p. 223. Lesser Antilles—Viecques, St. Thomas, Guadeloupe, Dominican<br />

Republic—Duchassaing and Michelotti, 1864, p. 41 (as Spongia rubens); St. Thomas,<br />

Tortole (?), St. Barthelemy—Duchassaing and Michelotti, 1864, p. 79 (as Amphimedon<br />

arborescens); Antilles—Schmidt, 1870, p. 37, (as Pachychalina rubens); Australia—South<br />

Australia—Carter, 1882, p. 276 (as Chalina rubens)}<br />

DISCUSSION. The species is attributed to Pallas whose red ramose sponge came from<br />

American Seas. De Laubenfels (1936a) has verified Duchassaing and Michelotti's record<br />

<strong>of</strong> H. rubens from their specimens. The type <strong>of</strong> their red Amphimedon arbor escens had<br />

apparently been lost. Cacochalina rubiginosa Schmidt (1870, p. 73) may be another<br />

synonym <strong>of</strong> H. rubens. Carter's Australian record, based on a light yellow sponge, is<br />

almost certainly erroneous.<br />

Haliclona erina de Laubenfels<br />

(Plate I, figs. 2, 3)<br />

Haliclona erina de Laubenfels, 1936b, p. 457 [type: Fort Randolph, Panama; USNM<br />

no. 22245]; idem, 1956, p. 3.<br />

[non] Haliclona erina, Burton, 1954, p. 223.<br />

HABITAT. Common in the turtle grass bed near Port Royal. It also occurs in the<br />

mangrove boat channel. Some <strong>of</strong> the mangrove specimens were growing over specimens<br />

<strong>of</strong> Geodia.<br />

SHAPE. A thickly encrusting to massive base, with volcano-shaped oscular projections.<br />

The sponge may also be somewhat ramose.<br />

COLOR. A dull dark green in life. The color is similar in specimens growing under


<strong>20</strong> A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

shaded and well-lit conditions. Preserved specimens become pale drab or dull yellowishgreen.<br />

CONSISTENCY. Firm. The sponge is very easily crumbled.<br />

SURFACE. Even, punctiform and smooth to the touch. The oscules are 3-4 mm in<br />

diameter. Most <strong>of</strong> the vents are apically placed on volcanic projections, 1-6 cm in height.<br />

The thick-walled projections enclose axial cloacal cavities which may be several centimeters<br />

long.<br />

ECTOSOME. No skeletal specialization. The scattered dermal pores are 30-60 ^ in<br />

maximum diameter. They pierce an aspiculous dermal membrane.<br />

ENDOSOME. Microcavernous. The skeleton is a spicular network with some spongin<br />

at the nodes. The network is composed <strong>of</strong> single spicules and ill-defined tracts with<br />

several rows <strong>of</strong> spicules. The meshes are typically 3-4 sided, with parallel sides 70-150 //,<br />

apart. Radial and connective bundles are well developed near the periphery, becoming<br />

as much as 90 jx in diameter. The tracts <strong>of</strong>ten project slightly beyond the fleshy surface.<br />

SPICULES. Robust oxeas, slightly curved to straight, 135-188 x 5-12 /x. A few long<br />

oxeas, apparently proper, are <strong>20</strong>0-270 ^ in length. The spicules narrow near the ends<br />

but are seldom strongly hastate. Mucronate spicules also occur. The developmental stages<br />

are thin and fusiform. A few styles, strongyles and centrotylote strongyles are present.<br />

Individual analyses, lengths in microns, 50 spicules each: 152-198; 163-188; 152-188; 141-<br />

176; 141-182; 135-182.<br />

DISTRIBUTION. Tropical Atlantic America: Atlantic coast <strong>of</strong> Panama—de Laubenfels,<br />

1936b, p. 457; Brazil—de Laubenfels, 1956, p. 3. American Atlantic Boreal: Newfoundland—Burton,<br />

1954, p. 223?<br />

DISCUSSION. TWO species <strong>of</strong> green haliclonids occur in the West Indies. Haliclona<br />

viridis (Duchassaing and Michelotti) de Laubenfels is said to have a limp, s<strong>of</strong>t consistency<br />

in life. Most <strong>of</strong> the reported specimens have spicules which are much less robust than<br />

those <strong>of</strong> my specimens. The North Carolina example <strong>of</strong> H. viridis reported by Wells<br />

et al. does have robust spicules. It differs in color and consistency from the Jamaican<br />

sponge. A Spicule slide prepared from de Laubenfels' Bahaman specimen at the American<br />

<strong>Museum</strong> <strong>of</strong> <strong>Natural</strong> <strong>History</strong> contains only thin fusiform oxeas. A sample <strong>of</strong> 50 spicules<br />

varied in length between 130-152 fi. The diameter was only 2-3 /x. A Bermudan specimen<br />

in the <strong>Yale</strong> <strong>Peabody</strong> <strong>Museum</strong>, identified by de Laubenfels, has oxeas, 123-145 x 2-3 JJL<br />

(25 spicules).<br />

De Laubenfels emphasized a brilliant green color and wide range <strong>of</strong> spicule size in<br />

his brief description <strong>of</strong> Haliclona erina. Oscular position was not described. Except for a<br />

less brilliant color, the Jamaican sponges are similar to H. erina and are provisionally<br />

considered to be conspecific with it.<br />

Burton's Newfoundland record is questionable in view <strong>of</strong> the locality and the inadequacy<br />

<strong>of</strong> the original description.<br />

Haliclona hogarthi n. sp.<br />

(Text-fig. 1; Plate II, fig. 1)<br />

Fig. 1. —Spicules <strong>of</strong> Haliclona hogarthi. Oxeas. YPM 5033. Holotype.


SYSTEMATICS 21<br />

HOLOTYPE. YPM 5033. Mangrove boat channel, July 13, 1961.<br />

HABITAT. Abundant on mangrove roots near Port Royal.<br />

SHAPE. The sponge consists <strong>of</strong> numerous, slender, anastomosing branches. The<br />

branches are <strong>of</strong>ten only several millimeters and are seldom more than 1 cm in diameter.<br />

COLOR. The living sponge has a light reddish-purple periphery. It gradually becomes<br />

drab toward the interior. In alcohol, the sponges are a pale to dark drab.<br />

CONSISTENCY. Limp, s<strong>of</strong>t, compressible, somewhat resilient.<br />

SURFACE. Punctiform. Even but wrinkled. The surface is microtuberculate as seen<br />

under a dissecting microscope at a magnification <strong>of</strong> 14 x. The numerous scattered oscules<br />

are 1-3 mm in diameter. They may be even with the surface or slightly raised.<br />

ECTOSOME. The dermal pores are 12-35 p, in diameter. They are separated by narrow<br />

aspiculous bands <strong>of</strong> flesh. At the surface many spicules are tangential while others project<br />

from the surface.<br />

ENDOSOME. An isodictyal, largely unispicular reticulation with 3- or 4-sided meshes.<br />

No spicule bundles are present. Spongin is present at nodes and occasionally envelops<br />

an entire spicule. The thin developmental stages <strong>of</strong> oxeas are scattered in the flesh.<br />

Aspiculous embryos are present in specimens collected in January, February and May.<br />

SPICULES. Oxeas, straight to slightly curved, hastate, occasionally mucronate, 117-157<br />

X 5-9 fi. A few styles occur.<br />

Individual analyses, lengths in microns, 50 spicules each: 128-152*; 123-140; 123-145;<br />

128-146; 117-152; 117-157.<br />

DISCUSSION. The species is characterized by its anastomosing form, reddish-purple<br />

color, limp consistency and hastate spicules. It is named in honor <strong>of</strong> Mr. W. E. Hogarth,<br />

<strong>of</strong> Kingston, Jamaica, whose encouragement is deeply appreciated.<br />

FAMILY DESMACIDONIDAE Gray, sensu de Laubenfels, 1936a<br />

GENUS DESMAPSAMMA Burton<br />

Desmapsamma Burton, (1934) has the spiculation <strong>of</strong> Desmacidon Bowerbank and a<br />

similar structure, aside from the presence <strong>of</strong> debris. De Laubenfels (1936a, p. 98) placed<br />

Desmapsamma in his heterogeneous family Psammascidae, which only obscures its relationships.<br />

Desmapsamma anchorata (Carter) Burton<br />

(Plate II, fig. 4)<br />

Fibularia anchorata Carter, 1882, p. 283 [type: Antigua; Brit. Mus. (Nat. Hist.)(?)]<br />

Desmapsamma anchorata, Burton, 1934, p. 547; idem, 1956, p. 131; idem, 1959, p. 239;<br />

Levi, 1959, p. 135.<br />

Desmacidon reptans Ridley and Dendy, 1886, p. 345 (fide Burton, 1934); idem, 1887, p.<br />

105; Lindgren, 1897, p. 482; idem, 1898, p. 303.<br />

Desmacidon carterianum Arndt, 1927, p. 147.<br />

HABITAT. Common on pilings at Port Royal and on coral heads in 10-12 feet <strong>of</strong> water<br />

near the cays.<br />

SHAPE. In a typical example a number <strong>of</strong> solid cylindrical branches arise from a<br />

common base. The branches may be several millimeters to more than a centimeter in<br />

diameter and <strong>of</strong>ten attain a height <strong>of</strong> 12 cm. Small oscules are sparsely scattered over<br />

their surface. In many specimens the base also gives rise to thick-walled oscular tubes.<br />

The tubes have an axial cloaca and wide apical oscules. The projections are typically<br />

1-2 cm in diameter and 1-3 cm in height. A few specimens have only the oscular type <strong>of</strong><br />

projection.<br />

COLOR. In life the exterior is a pale pink and the interior, a darker reddish-orange.<br />

In alcohol the sponges become beige to light gray but <strong>of</strong>ten retain a pinkish tinge.<br />

CONSISTENCY. Compressible, quite resilient.<br />

SURFACE. The surface is smooth to the touch but somewhat wrinkled. A dermal membrane<br />

can be detached with some difficulty. The oscules are 1-2 mm in diameter along


22 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

the branches and up to 5 mm in diameter on the oscular projections. All oscules have<br />

a membranous rim which is depressed with respect to the adjacent surface.<br />

ECTOSOME. The dermal membrane is supported by the plumose, slightly projecting<br />

ends <strong>of</strong> the interior spicule fibers. The dermal membrane contains sigmas, very numerous<br />

isochelas and scattered oxeas. Considerable amounts <strong>of</strong> sand and spicule fragments are<br />

scattered in the skin <strong>of</strong> all the examined specimens. The proper megascleres and debris<br />

form a reticulate pattern when they are condensed in inter-ostial partitions. The dermal<br />

pores are 30-65 /x in diameter.<br />

ENDOSOME. The interior contains scattered oxeas and an irregular reticulation <strong>of</strong><br />

spicule tracts. The latter are <strong>of</strong>ten 10-90 p in diameter and 100 ^ apart. Longitudinal<br />

strands are well developed. They give <strong>of</strong>f radial bundles which end in dermal spicule<br />

tufts. Microscleres <strong>of</strong> both types are common in the flesh. Very little foreign material is<br />

present in the interior.<br />

SPICULES, (a) Oxeas, slightly curved to straight, mostly hastate, 140-<strong>20</strong>0 X 3-7 p. The<br />

oxeas <strong>of</strong> the interior spicule tracts are stout. Most <strong>of</strong> the dermal and interstitial megascleres<br />

are thin, (b) Sigmas, 10-43 X 1-2 fi. (c) Arcuate isochelas, 10-23 ^ in length, with<br />

a shaft diameter <strong>of</strong> 1-2 /x. The toothed ends are as much as 7 fi in diameter. The ends<br />

are 4 jn long in a spicule <strong>of</strong> 12 p total length, and 7/J, long in one <strong>of</strong> 17 fi total length.<br />

Individual analyses, lengths in microns:<br />

Oxeas Sigmas Isochelas<br />

145-195 (50) 12-36 (25) 10-18 (25)<br />

145-188 (50) 10-37 (25) 10-18 (25)<br />

140-199 (50) 13-40 (25) 10-22 (50)<br />

143-<strong>20</strong>0 (50) 12-43 (50) 12-23 (50)<br />

DISTRIBUTION. Tropical Atlantic America: Brazil—Ridley and Dendy, 1886, p. 345<br />

and 1887, p. 105 (as Desmacidon reptans); Antigua—Carter, 1882, p. 283 (as Fibularia<br />

anchorata); Curasao—Arndt, 1927, p. 147 (as Desmacidon carterianum); Tropical West<br />

Africa—Burton, 1956, p. 131, and LeVi, 1959, p. 135. Indo-Pacific: Zanzibar, East Africa—<br />

Burton, 1959, p. 239; China Sea—Lindgren, 1897, p. 482 and 1898, p. 303 (as Desmacidon<br />

reptans); Great Barrier Reef, Australia—Burton, 1934, p. 547.<br />

DISCUSSION. The Jamaican specimens are most similar in structure to the Brazilian<br />

sponges described as Desmacidon reptans by Ridley and Dendy. My specimens differ only<br />

in having better developed spicule tracts in the interior. The West Indian specimens <strong>of</strong><br />

Arndt and Carter had considerably more debris, with foreign material abundant in the<br />

interior. It should be noted that all <strong>of</strong> my specimens were growing on wood or coral<br />

surfaces, with little sand available. The specimens <strong>of</strong> Lindgren and LeVi differ from<br />

mine in having dermal oxeas slightly smaller in length than those <strong>of</strong> the interior.<br />

The specimens attributed to the species, although from widely separated localities,<br />

are very similar in structure. Burton (1934, p. 547) examined the specimens <strong>of</strong> Ridley<br />

and Dendy, Carter and Lindgren and concluded that they were conspecific.<br />

NEOFIBULARIA n. gen.<br />

A reticulate skeleton without dermal specialization. The megascleres are strongyles.<br />

The microsclere complement includes raphides and <strong>of</strong>ten sigmas. Fibularia massa Carter<br />

(1882) is the type-species.<br />

Ne<strong>of</strong>ibularia is proposed as a new name for Fibularia Carter. The latter is preoccupied<br />

for an echinoderm, as noted by Vosmaer (1886, p. 392, 400) and de Laubenfels<br />

(1936a, p. 51). Carter (1882) included three West Indian species in his genus which was<br />

undefined. Fibularia massa had strongyles, raphides and sigmas. F. ramosa had oxeas and<br />

sigmas, and was transferred to Gelliodes by de Laubenfels (1936a, p. 53). F. anchorata,<br />

with oxeas, sigmas and foreign material, was made the type species <strong>of</strong> Desmapsamma by<br />

Burton (1934, p. 547).<br />

In 1886 Carter described an Australian sponge with oxeas and sigmas as Fibulia<br />

carnosa. Fibulia was not given a diagnosis. De Laubenfels (1936a, p. 51) considered


SYSTEMATICS 23<br />

Fibulia to be a direct substitution for the preoccupied Fibularia. Burton (1929, p. 142)<br />

had already established a new genus Plumocolumella for Fibulia carnosa on the assumption<br />

that Fibulia was a misprint for Fibularia. Burton later concluded (1936b, p. 142)<br />

that "no substantial evidence <strong>of</strong> a misprint" was available. He therefore considered<br />

Fibulia to be a valid genus with F. carnosa Carter as its type-species. Carter gave no<br />

indication that Fibulia was to be a substitute for Fibularia. A new generic name is<br />

therefore necessary for Fibularia massa Carter.<br />

In addition to the type species, Ne<strong>of</strong>ibularia includes the following species:<br />

Fibulia nolitangere (Duchassaing and Michelotti, 1864, p. 82) de Laubenfels, 1936a, p. 51.<br />

Fibulia raphidifera (Topsent, 1889, p. 16) de Laubenfels, 1950a, p. 78.<br />

Fibulia bermuda de Laubenfels, 1950a, p. 52.<br />

Ne<strong>of</strong>ibularia massa (Carter) new combination<br />

Fibularia massa Carter, 1882, p. 282 [type: Bahamas; Brit. Mus. (Nat. Hist.) (?)]<br />

Gellius massa, Arndt, 1927, p. 151.<br />

Fibulia massa, de Laubenfels, 1950a, p. 53; idem, 1953a, p. 527.<br />

HABITAT. A common species near the cays. It <strong>of</strong>ten grows on the underside <strong>of</strong> projecting<br />

ledges in shallow water. In deeper water (10-15 feet) it grows on coral masses.<br />

SHAPE. The shallow water specimens are laterally spreading masses several centimeters<br />

in thickness. Their large cloacae lead to terminal openings which are flush with the<br />

surface. The <strong>of</strong>fshore specimens may be vasiform or consist <strong>of</strong> several thick-walled hollow<br />

projections arising from a basal mass. Large vases <strong>of</strong>ten reach 12 cm in height. Their<br />

walls reach a thickness <strong>of</strong> 2-4 cm and their cloacal openings, a diameter <strong>of</strong> 2-4 cm.<br />

COLOR. In life the peripheral 1-2 mm are a reddish or mustard brown. The color<br />

gradually fades to a dull yellow toward the interior. Preserved specimens are beige to<br />

drab with a greenish tinge and occasionally have purple tinges.<br />

CONSISTENCY. S<strong>of</strong>t, very easily crumbled.<br />

SURFACE. The outer and cloacal surfaces are fibrohispid. The outer surface is rough<br />

to the touch but even. It is pierced by a few oscular (?) openings which are less than 0.5<br />

mm in diameter. Numerous exhalant canals, 0.5-5 mm in diameter, empty into the<br />

cloacal cavity. They are <strong>of</strong>ten continued upward as longitudinal grooves in the cloacal<br />

surface.<br />

ECTOSOME. Fibrohispid. No dermal specialization.<br />

ENDOSOME. Cavernous. The skeleton is a coarse-meshed, symmetrical reticulation <strong>of</strong><br />

stout spicule tracts, which are 40-150 //, in diameter. The rectangular meshes are 300-<br />

1000 ji in maximum width. Raphides, sigmas and some megascleres are strewn in the<br />

flesh.<br />

SPICULES, (a) Strongyles, slightly to strongly curved, 261-386 X 5-10 /x. (b) Raphides,<br />

68-1 50JU, in length. They occasionally reach 3 p in diameter but are usually much thinner.<br />

Many are grouped into trichodragmata, 30-60 ^ in diameter, (c) Sigmas, 16-26 /x.<br />

Individual analyses, length in microns:<br />

Strongyles Raphides Sigmas<br />

261-334 (50) 68-122 (25) 17-24 (25)<br />

281-386 (50) 79-150 (25) 18-26 (25)<br />

292-357 (50) 72-133 (25) 18-23 (25)<br />

26^358 (50) 76-130 (25) 16-23 (25)<br />

DISTRIBUTION. Tropical Atlantic America: Florida—de Laubenfels, 1950a, p. 53 (as<br />

Fibulia massa); West coast, Florida—de Laubenfels, 1953a, p. 527 (as Fibulia massa);<br />

Bahamas—Carter, 1882, p. 282 (as Fibularia massa); Curasao—Arndt, 1927, p. 151 (as<br />

Gellius massa).<br />

DISCUSSION. A graded series was observed between vasiform and thickly incrusting<br />

specimens. Carter's sponge was <strong>of</strong> the latter type. The species is very irritating to the skin.


24 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

GENUS IOTROCHOTA Ridley<br />

Iotrochota birotulata (Higgin) Ridley<br />

Halichondria birotulata Higgin, 1877, p. 296 [type: Venezuela; Brit. Mus. (Nat. Hist.) (?)]<br />

[non] Halichondria birotulata, Carter, 1886, p. 52 (fide Dendy, 1896); idem, 1887, p. 72.<br />

Iotrochota birotulata, Ridley, 1884, p. 433; de Laubenfels, 1932a, p. 37; idem, 1936a, p.<br />

49; idem, 1953a, p. 522.<br />

HABITAT. Common on pilings at Port Royal and on coral masses <strong>of</strong>fshore from the<br />

cays.<br />

SHAPE. Typically ramose with branches 1-2 centimeters in diameter. One massive<br />

specimen was collected.<br />

COLOR. Very dark purple, nearly black, with a greenish surface sheen in life. A<br />

purplish slime is emitted when the sponge is handled. Dried and preserved specimens<br />

are purplish black.<br />

CONSISTENCY. Tough, rather stiff.<br />

SURFACE. Numerous conules and connecting ridges make the surface very uneven.<br />

The narrow sharp-pointed conules are 1-3 mm in height and 1-8 mm apart. The shallow<br />

depressions between the elevated areas are covered by a thin skin. The oscules are small,<br />

scattered and infrequent. Orange zoanthids are present on the surface.<br />

ECTOSOME. The skin contains sparsely scattered, long, thin styles and numerous<br />

birotules. It is pierced by numerous ostia which are <strong>20</strong>-30 fi in diameter. In places, the<br />

surface is underlain by subdermal cavities which are <strong>of</strong>ten <strong>20</strong>0 /x in depth.<br />

ENDOSOME. The skeleton includes a reticulation <strong>of</strong> spicule bundles and scattered<br />

interstitial megascleres. Birotules are most abundant near the canals and periphery<br />

The spicule tracts are 35-180 p in diameter and 170-650 ^ apart. Stout bundles extend<br />

into the conules. No localization <strong>of</strong> megasclere types could be detected in the endosome.<br />

A specimen collected during November, 1960, contains numerous densely pigmented<br />

aspiculous embryos, as large as 500 /x in longest diameter.<br />

SPICULES, (a) Strongyles, mostly curved, 117-286 p, in length. The diameter varies between<br />

3-8 /x, with the longest spicules typically thin and occasionally anisostrongylote.<br />

(b) Styles, less frequent, mostly curved, 129-253 x 5-10 /*. The longer spicules, such as<br />

those in the ectosome, are frequently thin, (c) Oxeas, hastate, rare or absent, 152-188 p.<br />

Their diameter is similar to that <strong>of</strong> the styles and strongyles. (d) Birotules, 10-18 /JL in<br />

length.<br />

Individual analyses, lengths in microns:<br />

Strongyles<br />

Styles<br />

Oxeas Birotules<br />

117-217 (50)<br />

140-246 (50)<br />

117-258 (50)<br />

152-286 (50)<br />

present<br />

- present<br />

present<br />

130-182 (50)<br />

159-253 (30)<br />

129-193 (30)<br />

130-186 (30)<br />

present<br />

present<br />

present<br />

152-188 (25)<br />

183 (1)<br />

none<br />

163 (2)<br />

rare<br />

rare<br />

rare<br />

13-16 (25)<br />

13-17 (25)<br />

12-14 (25)<br />

12-15 (25)<br />

13-18 (25)<br />

13-17 (25)<br />

10-13 (25)<br />

DISTRIBUTION. Tropical Atlantic America: Dry Tortugas, Florida—de Laubenfels,<br />

1932a, p. 37, 1936a, p. 49; West coast, Florida—de Laubenfels, 1953a, p. 522; Venezuela—<br />

Higgin, 1877, p. 296 (as Halichondria birotulata); Bahamas—Higgin, 1877, p. 296 (as<br />

Halichondria birotulata); Jamaica—Higgin, 1877, p. 296 (as Halichondria birotulata),<br />

de Laubenfels, 1932a, p. 37; Indo-Pacific: Mergui Archipelago, <strong>of</strong>f Burma—Carter, 1887,<br />

p. 72 (as Halichondria birotulata)(?).<br />

DISCUSSION. Oxeas are known to occur in the genus, being common, for example, in<br />

I. acerata Dendy (1896). They have not been recorded previously for 7. birotulata. Although<br />

rare at best, their uniformity <strong>of</strong> shape indicates that they are proper to the<br />

sponge.<br />

De Laubenfels reported that the birotules may be rare or absent. Higgin's sponge


SYSTEMATICS 25<br />

from the Bahamas was described as having no microscleres. Numerous birotules were<br />

present in the Jamaican specimens examined by Higgin, de Laubenfels and myself.<br />

Spongin could not be detected in the present specimens. Higgin found it to be variable<br />

in quantity and least abundant in his Jamaican specimen.<br />

Dendy (1896) re-examined Carter's 1886 Australian sponge and found it to be conspecific<br />

with Iotrochota coccinea (Carter, 1887) rather than with Higgin's species. Carter's<br />

1887 record <strong>of</strong> an Indian Ocean /. birotulata, based on an undescribed reddish-purple<br />

specimen, must be questioned.<br />

GENUS GELLIODES Ridley<br />

Gelliodes areolata (Wilson) new combination<br />

(Plate II, fig. 3)<br />

Pachychalina areolata Wilson, 1902a, p. 392 [type: <strong>of</strong>f Vieques; USNM no. 7671.]<br />

Haliclona areolata, de Laubenfels, 1936a, p. 39; Wells et al., 1960, p. <strong>20</strong>7.<br />

Haliclona excelsa, de Laubenfels (not Schmidt, 1870), 1947, p. 36 (fide Wells et al., 1960.)<br />

HABITAT. Abundant on pilings at Port Royal.<br />

SHAPE. Solid, cylindrical. The sponge may consist <strong>of</strong> a single flattened cylinder or a<br />

cluster <strong>of</strong> cylindrical branches. The branches are typically 2-3 cm in diameter. Many<br />

specimens are 10-<strong>20</strong> cm in maximum length.<br />

COLOR. In life, the sponges are a dull reddish pink or a dull purple. They become<br />

beige in alcohol.<br />

CONSISTENCY. Compressible, resilient.<br />

SURFACE. Rough to the touch but macroscopically even. The scattered oscules are<br />

1.5-3 mm in diameter. They are <strong>of</strong>ten raised slightly above the surface. Numerous<br />

zoanthids are usually present on the surface.<br />

ECTOSOME. NO dermal specialization. The internal fibers subdivide repeatedly near<br />

the surface where they end in non-plumose tufts.<br />

ENDOSOME. A coarse fibroreticulation with rectangular meshes, 100-350 /x in width.<br />

The fibers are 25-235 ^ in diameter. They consist <strong>of</strong> numerous, closely packed rows <strong>of</strong><br />

spicules cemented by a varying amount <strong>of</strong> spongin. Some megascleres are scattered in the<br />

flesh. Microscleres were absent in five analyzed specimens but abundant in the sixth.<br />

SPICULES, (a) Robust oxeas, nearly straight or slightly curved, 182-276 x 5-18 p. The<br />

abruptly narrowed ends are sometimes rounded or irregular in outline. A varied number<br />

<strong>of</strong> megascleres are really styles or strongyles. The latter are <strong>of</strong>ten only 150 /n in length.<br />

A few abnormal spicules in all six specimens have a short, oblique spine or ray near<br />

one end. (b) Sigmas, in one specimen, 13-17 X 1 ft-<br />

Individual analyses, length in microns:<br />

Oxeas (50) Styles (25)<br />

182-258 176-234<br />

<strong>20</strong>0-258 182-234<br />

211-269 182-246<br />

199-264 194-234<br />

211-252 182-234<br />

217-276 present<br />

Strongyles (25)<br />

188-217<br />

152-264<br />

146-211<br />

164-176<br />

176-234<br />

present<br />

Sigmas<br />

none<br />

none<br />

none<br />

none<br />

none<br />

13-17 (,25)<br />

DISTRIBUTION. Tropical Atlantic America: North Carolina—de Laubenfels, 1947, p. 37<br />

(as Haliclona excelsa), Wells et al., 1960, p. <strong>20</strong>7; Puerto Rico—Wilson, 1902a, p. 392 (as<br />

Pachychalina areolata).<br />

DISCUSSION. In form, architecture and oxea size the Jamaican sponge is very similar<br />

to Pachychalina areolata Wilson (1902a, p. 392) <strong>of</strong> Puerto Rico. Wilson found no sigmas<br />

in his species, and I have found none in a slide prepared from his type. They were<br />

lacking in 5 or 6 Jamaican sponges. Puerto Rican sponges in the <strong>Yale</strong> <strong>Peabody</strong> <strong>Museum</strong>,<br />

which are certainly conspecific with the Jamaican ones, do have a varied number <strong>of</strong> thin<br />

sigmas. (W. D. Hartman, personal communication). Wells et al. (1960, p. <strong>20</strong>7) have


26 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

attributed a North Carolina sponge, lacking microscleres, to Wilson's species. Their<br />

specimen is also similar to the Jamaican one in architecture and megasclere size. Wells<br />

et al. list Pachychalina millepora Verrill (1907, p. 336) as a synonym. Verrill's species and<br />

the Puerto Rican Pachychalina aurantiaca (Lendenfeld) <strong>of</strong> Wilson (1902a, p. 393) are<br />

certainly closely related to, if not conspecific with, the present species.<br />

FAMILY ADOCIIDAE de Laubenfels, 1934<br />

A simple megasclere complement, with diactinal spicules. Microscleres are <strong>of</strong>ten<br />

present and may include sigmas, chelas, toxas and raphides. A distinct tangential dermal<br />

skeleton is present. The architecture is reticulate.<br />

De Laubenfels (1936a) placed the Adociidae in the order Poecilosclerida despite their<br />

lack <strong>of</strong> spicule specialization. De Laubenfels himself had pointed out the close relationship<br />

<strong>of</strong> the adociids with the haplosclerid sponges. Many adociid genera can be distinguished<br />

from haliclonid or desmacidonid genera only by the greater development <strong>of</strong><br />

a dermal skeleton. Species with an intermediate type <strong>of</strong> architecture, such as the Jamaican<br />

Sigmadocia caerulea, are known in many such cases. The Adociidae, indeed, are probably<br />

a heterogeneous group. They can be provisionally maintained as a family, at least for<br />

convenience, within the order Haplosclerida. Levi (1958) has also commented on the<br />

similarities between the adociids and haplosclerids <strong>of</strong> de Laubenfels* classification.<br />

GENUS ADOCIA Gray<br />

Adocia carbonaria (Lamarck) new combination<br />

(Plate III, figs. 2, 3)<br />

Spongia carbonaria Lamarck, 1813, p. 375 (type: American seas; Mus. nat. Hist. nat.<br />

Paris.)<br />

Thalysias carbonaria, Duchassaing and Michelotti, 1864, p. 83.<br />

Pachychalina carbonaria, Arndt, 1927, p. 152.<br />

Phloeodictyon carbonaria, Topsent, 1930, p. 26.<br />

[non] Pellina carbonaria, de Laubenfels, 1936a, p. 68; idem, 1954a, p. 102; idem, 1955,<br />

p. 139.<br />

Thalysias proxima Duchassaing and Michelotti, 1864, p. 84 (fide Arndt, 1927.)<br />

HABITAT. The species is common in the turtle grass bed at Drunkenman's Cay. It also<br />

occurs in a similar habitat at Dunn's River on the North Coast <strong>of</strong> Jamaica.<br />

SHAPE. Thick incrustations or repent branches. The incrusting examples are 2-3 cm<br />

in thickness. The branches, always solid, are 0.5-1.5 cm in diameter.<br />

COLOR. The living sponges are entirely black. When dried or preserved they become<br />

a very dark brown. Alcohol becomes blue-black when the sponge is placed in it.<br />

CONSISTENCY. Brittle and easily crumbled even when alive.<br />

SURFACE. Smooth to the touch, but uneven. The oscules are 1-3 mm in diameter. They<br />

are scattered over the surface in both incrusting and branching specimens. Many oscules<br />

are the apertures <strong>of</strong> exhalant canals which extend straight downwards into the sponge.<br />

The oscules may be flush with the surface, or raised on low volcano-shaped elevations.<br />

Although the skin is easily detachable, macroscopic subdermal spaces are not present.<br />

ECTOSOME. The dermal membrane is supported by a three- to four-sided spicule network.<br />

The meshes are generally unispicular but occasionally have two or three spicules<br />

on a side. The interstices contain one to several dermal pores which are 30-75 p in diameter.<br />

Small quantities <strong>of</strong> spongin are present at the nodes* Spicules sometimes join a<br />

node at their mid-length. Numerous pigment cells, 7 p in diameter, are present in both<br />

the ectosome and interior.<br />

ENDOSOME. The interior has a largely unispicular network, with some but not all <strong>of</strong><br />

the spicules bound by spongin nodes. The skeleton also includes an irregular reticulation<br />

<strong>of</strong> spicule tracts, which are 30-45 p in diameter.<br />

SPICULES. Oxeas, mostly curved, 176 x 246 X 3-9 p. The ends may be rounded, mucronate,<br />

or slightly stair-stepped. The stair-stepped spicules may have truncate ends. A few<br />

styles and strongyles are present. The developmental stages are curved, fusiform oxeas.


SYSTEMATICS 27<br />

Individual analyses, lengths in microns, 50 spicules each: Oxeas, 199-229; 176-246;<br />

188-234.<br />

DISTRIBUTION. Tropical Atlantic America: Lamarck, 1813, p. 375 (as Spongia carbonaria);<br />

Lesser Antilles—Guadeloupe, St. Croix, St. Barthelemy, St. Thomas—Duchassaing<br />

and Michelotti, 1864, p. 83 (as Thalysias carbonaria); St. Thomas—Duchassaing and<br />

Michelotti, 1864, p. 84 (as Thalysias proximo); Curasao—Arndt, 1927, p. 152 (as Pachychalina<br />

carbonaria).<br />

DISCUSSION. In spicule shape the present specimens are somewhat similar to the black<br />

Pellina carbonaria studied by de Laubenfels (1936a, p. 66) in the Dry Tortugas. That<br />

sponge <strong>of</strong>ten grew in sand, with only its long oscular projections protruding above the<br />

surface. He considered it to be conspecific with Spongia carbonaria Lamarck but later<br />

(1954a, p. 107) expressed some reservations about the identification. The spicules <strong>of</strong> the<br />

Dry Tortugas sponges are smaller than those <strong>of</strong> either the Jamaican sponge or <strong>of</strong> Lamarck's<br />

specimen. In addition the spicule was described as "fundamentally a strongyle,"<br />

with the ends "sometimes provided with caps <strong>of</strong> successively smaller diameter."<br />

De Laubenfels' sponge differs further from the present specimen in having a dermal<br />

skeleton <strong>of</strong> irregularly strewn spicules and subdermal spaces. It was described as being<br />

very dark green rather than black. The consistency was said to be delicately elastic and<br />

easily torn, rather than stiff. De Laubenfels, who paid close attention to color changes<br />

in alcohol, made no mention <strong>of</strong> any blue-black exudate. The oscules were clearly <strong>of</strong> the<br />

Pellina type. It could be that the delicate projections were suppressed in the Jamaican<br />

sponges, which were indeed subject to considerable surf. The presence <strong>of</strong> delicate, stiff,<br />

slender branches in the Jamaican specimens weighs against such a conclusion. The<br />

numerous differences strongly suggest that the Dry Tortugas sponge is not conspecific<br />

with the Jamaican.<br />

Pellina carbonaria (Lamarck) as reported from the central Pacific by de Laubenfels<br />

(1954a, p. 102) does have large spicules but also has oscular chimneys and lacks a blueblack<br />

exudate. Its relationship to the sponges <strong>of</strong> the Dry Tortugas and Jamaican is<br />

uncertain.<br />

The poorly preserved type <strong>of</strong> Spongia carbonaria Lamarck has been restudied by<br />

Topsent (1930). It bore no trace <strong>of</strong> oscular projections. Its curious system <strong>of</strong> cavities and<br />

lamellae may be the result <strong>of</strong> drying. The oxeas had abrupt but not stair-stepped points.<br />

They were similar in length to those <strong>of</strong> the Jamaican specimens. Duchassaing and<br />

Michelotti's Thalysias carbonaria resembles the Jamaican specimens in shape and color.<br />

No information is available on its spiculation. Arndt's sponges from Curasao are probably<br />

conspecific with the Jamaican ones. His specimens had a similar shape, color,<br />

dermal skeleton and spicule size range. The tracts <strong>of</strong> the main skeleton were somewhat<br />

better developed. The spicules <strong>of</strong>ten had wedge-shaped points but apparently were not<br />

stair-stepped.<br />

The Jamaican specimens are somewhat doubtfully considered to be conspecific with<br />

those <strong>of</strong> Lamarck, Arndt, and Duchassaing and Michelotti. The lack <strong>of</strong> oscular tubes<br />

places the species in the genus Adocia.<br />

Adocia implexiformis n. sp.<br />

(Text-fig. 2; Plate II, fig. 2)<br />

Fig. 2. —Spicules <strong>of</strong> Adocia implexiformis. Oxeas. YPM 5034. Holotype.<br />

HOLOTYPE. YPM 5034, Mangrove boat channel, Jan. 25, 1961.<br />

HABITAT. Abundant on mangrove roots.


28 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

SHAPE. Thick-walled, cylindrical oscular projections arise from the base. They are<br />

typically 2-4 cm in height and 1-2 cm in diameter. Adjacent oscular lobes may be fused<br />

laterally. Slender, solid projections, 0.5-1 cm in height, and 2-3 mm in diameter, are<br />

<strong>of</strong>ten present on distal parts <strong>of</strong> the sponge.<br />

COLOR. The living sponge is a dull purple, devoid <strong>of</strong> any reddish tinge. The sponges<br />

are drab in alcohol.<br />

CONSISTENCY. The sponge retains its shape when removed from water. It is compressible<br />

but very easily crumbled.<br />

SURFACE. Even. The oscules are 2-10 mm in diameter. They are the openings <strong>of</strong><br />

elongate axial cavities in the oscular lobes.<br />

ECTOSOME. A detachable dermal membrane is present. It is supported by a tangential,<br />

isodictyal, largely unispicular network <strong>of</strong> oxeas. Spongin is present at the nodes. The<br />

three- to four-sided meshes contain closely-spaced ostia which are 12-35 ^ in diameter. A<br />

few large subdermal spaces are present.<br />

ENDOSOME. A spicular network similar to that <strong>of</strong> the ectosome. The meshes occasionally<br />

have 2 or 3 rows <strong>of</strong> spicules on a side but long tracts are absent.<br />

SPICULES. Oxeas, straight to slightly curved, 129-182 X 5-10 /*. Thin, sharp-pointed<br />

developmental stages are present. The mature spicules have blunt ends, and a few are<br />

styles and strongyles. The ends <strong>of</strong> many oxeas are narrow, but rounded, rather than<br />

sharp-pointed. Individual analyses, lengths in microns, 50 spicules each: 145-174; 145-<br />

182*; 137-181; 129-179.<br />

DISCUSSION. In form, the present species resembles the heterogeneous assemblage <strong>of</strong><br />

sponges attributed to Haliclona implexa (Schmidt). The sponges assigned to the latter<br />

species lack a special dermal skeleton. A. chilensis (Thiele, 1905) <strong>of</strong> Chile also has oscular<br />

projections. It differs from the Jamaican species in having strongly strongylote spicules,<br />

spicule tracts and well-developed subdermal cavities. A. grossa (Schmidt), as recorded by<br />

Topsent (1925a) from Naples, is similar in form and spicule range to the present species.<br />

It differs in having an uneven surface and multispicular tracts. The Philippine projection-bearing<br />

A. baeri (Wilson, 1925) de Laubenfels, 1935b, has a tuberculate surface, dark<br />

color and small oxeas. A. tub if era (George and Wilson, 1919) Hartman, 1958, <strong>of</strong> North<br />

Carolina has small oscular projections on anastomosing branches. The skeleton includes<br />

spongin fibers.<br />

Adocia albifragilis n. sp.<br />

(Text-fig. 3)<br />

Fig. 3. —Spicules <strong>of</strong> Adocia albifragilis. Oxeas. YPM 5035. Holotype.<br />

HOLOTYPE. YPM 5035. Drunkenman's Cay, June 24, 1961.<br />

HABITAT. Under rocks at the cays, in a few feet <strong>of</strong> water.<br />

SHAPE. Thin crusts, reaching a maximum thickness <strong>of</strong> 2-3 mm.<br />

COLOR. The living sponge is white. It is pale, nearly white in alcohol.<br />

CONSISTENCY. Compressible, but easily torn.<br />

SURFACE. Even. Under a magnification <strong>of</strong> 14 x, the surface appears micro tuberculate<br />

but not hispid. The oscules are about 2 mm in diameter and flush with the surface.<br />

ECTOSOME. The surface has a tangential network closely connected to the skeleton <strong>of</strong><br />

the interior. No large subdermal spaces are present. The sides <strong>of</strong> the skeletal network<br />

may consist <strong>of</strong> single spicules or several loosely arranged spicules. Compact tracts do not<br />

occur. Small amounts <strong>of</strong> spongin are present at the nodes. A node may include as many<br />

50//<br />

I


SYSTEMATICS 29<br />

as 10-12 spicules. The meshes are thus <strong>of</strong>ten much less than a spicule length in maximum<br />

width. Only a few dermal pores, <strong>20</strong>-40 ^ in diameter, were detected.<br />

ENDOSOME. The internal skeleton is similar to that <strong>of</strong> the dermal layer. Most, if not<br />

all, <strong>of</strong> the spicules are bound into the network.<br />

SPICULES. Oxeas, thin, straight to slightly curved, 145-174 x 3-5 p. The points may<br />

be fusiform, hastate or mucronate. Many spicules are narrow but rounded at the ends.<br />

A small minority <strong>of</strong> spicules are styles or strongyles. Their rounded ends are frequently<br />

narrow. Individual analyses, length in microns, 50 spicules each: 145-174; 145-163*.<br />

DISCUSSION. The sponge is characterized by the presence <strong>of</strong> long thin spicules which<br />

are very <strong>of</strong>ten blunt. The frequent occurrence <strong>of</strong> nodes with many spicules is also noteworthy.<br />

Adocia neens (Topsent, 1918), reported for the West Indies by de Laubenfels (1936a),<br />

also has thin blunt spicules. Its spicules are smaller than in my specimen and are mostly<br />

strongylote. The dermis in A. neens has a conspicuous protoplasmic as well as spicular<br />

network.<br />

GENUS PELLINA Schmidt<br />

Pellina coeliformis n. sp.<br />

(Text-fig. 4; plate III, fig. 1)<br />

Fig. 4. —Spicules <strong>of</strong> Pellina coeliformis. Oxeas. YPM 5036. Holotype.<br />

HOLOTYPE. YPM 5036. Mangrove boat channel, July 30, 1959.<br />

HABITAT. The species is abundant in the Kingston Port Royal boat channel. It frequently<br />

grows on the surface <strong>of</strong> Geodia gibberosa.<br />

SHAPE. The sponge consists <strong>of</strong> an incrusting base, several millimeters in thickness,<br />

from which arise paper-thin oscular tubes. The tubes are usually 3-5 cm in height and<br />

several millimeters in diameter. A few specimens have wider oscular projections. The<br />

tubes occasionally branch near the base.<br />

COLOR. In life, the basal region is a pale <strong>of</strong>f-white color, while the tubes are white.<br />

In alcohol the base becomes beige and the tubes semi-transparent.<br />

CONSISTENCY. The base is slightly compressible. The tubes are bendable but easily torn.<br />

SURFACE. Smooth, even. Longitudinal tracts are visible to the unaided eye in the tube<br />

walls.<br />

ECTOSOME. Little dermal specialization. The outermost layer is a tangential, largely<br />

unispicular, three- or four-sided network. Small quantities <strong>of</strong> spongin are present at the<br />

nodes. The dermal pores are 15-35 ^ in diameter. The dermal layer grades into the main<br />

skeleton, and can be detached only with difficulty. No large subdermal spaces are present.<br />

ENDOSOME. The tubes are supported by numerous longitudinal spicule bundles, 30-<br />

80 n> in diameter, which occasionally branch and anastomose. Adjacent bundles are<br />

75-<strong>20</strong>0 JJL apart. They are crossed by connective fibers, <strong>20</strong>-35 fx in diameter, which are<br />

<strong>of</strong>ten obliquely oriented. The internal skeleton also includes a unispicular network,<br />

similar to that <strong>of</strong> the surface. The basal skeleton has a less regular arrangement. It includes<br />

spicule bundles, scattered spicules and, in places, a unispicular network.<br />

SPICULES. Oxeas, usually slightly curved and rather abruptly pointed, 117-158 X 5-7 /x.<br />

Less robust developmental forms are also present. Individual analyses, lengths in microns,<br />

50 spicules each: 123-158; 117-146; 123-152; 117-152; 117-146; 117-146; 123-145*.<br />

DISCUSSION. TWO species <strong>of</strong> Pellina have been recorded previously from the West


30 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

Indies. P. carbonaria (Lamarck) as described by de Laubenfels from Dry Tortugas is a<br />

black sponge with larger oxeas and a network <strong>of</strong> subdermal canals. The white P. coela<br />

de Laubenfels (1950a) <strong>of</strong> Bermuda differs from the Jamaican sponges by its smaller<br />

spicules and less robust spicule tracts. The type species <strong>of</strong> Pellina, P. semitubulosa, described<br />

from the Mediterranean, is also white in color. The Adriatic specimens studied<br />

by Babic (1922) and Keller (1878) do have spicules similar in size to those <strong>of</strong> my specimens.<br />

Topsent's Mediterranean specimen, however, has more robust oxeas and a second<br />

category <strong>of</strong> small ectosomal microxeas. He reported an extensive network <strong>of</strong> subdermal<br />

canals in the basal regions <strong>of</strong> his material. Schmidt (1862, 1870) and Topsent both<br />

emphasized the presence <strong>of</strong> a well-developed, easily detachable skin in their specimens.<br />

The Jamaican sponge, by contrast, has a closely adherent dermal skeleton.<br />

A<br />

GENUS SIGMADOCIA de Laubenfels<br />

Sigmadocia caerulea n. sp.<br />

(Text-fig. 5; plate III, fig. 4)<br />

50//<br />

Fig. 5. —Spicules <strong>of</strong> Sigmadocia caerulea. A. Oxea. B. Sigmas. YPM 5037. Holotype.<br />

HOLOTYPE. YPM 5037. Rasta's wreck. July 19, 1959.<br />

HABITAT. Common on pilings and mangrove roots on the outside <strong>of</strong> the Port Royal<br />

mangrove thicket. Sponges <strong>of</strong> similar appearance were collected at Drunkenman's Cay in<br />

a sandy turtle grass bed.<br />

SHAPE. The base is thickly incrusting to massive. Many <strong>of</strong> the oscules, particularly in<br />

the Port Royal specimens, are at the apex <strong>of</strong> cylindrical to volcano-shaped projections<br />

ivhich may be several centimeters in height. The projections have axial cloacae and thick<br />

-walls.<br />

COLOR. Both the surface and the interior are light blue in the living sponge. The<br />

specimens become beige to drab in alcohol.<br />

CONSISTENCY. The Port Royal specimens are s<strong>of</strong>t but very easily broken. The specimens<br />

from Drunkenman's Cay are heavily penetrated by coarse algal strands. They are<br />

stiff to somewhat compressible.<br />

SURFACE. The oscules may be as much as 5 mm in diameter. The surface, while even,<br />

varies from smooth to rough. Some areas <strong>of</strong> the sponge, particularly on the base, have<br />

a detachable dermal layer.<br />

ECTOSOME. Parts <strong>of</strong> the sponge are covered by a tangential, largely unispicular network.<br />

The three- or four-sided network is bound by small amounts <strong>of</strong> spongin at the<br />

nodes. Several dermal pores may be inclosed within a mesh. Typical pore sizes are<br />

<strong>20</strong> X 17, 12 X <strong>20</strong>, and 27 X 24 ^. Small subdermal spaces may be present. In other<br />

regions, the architecture is <strong>of</strong> the haliclonid type, with spicules projecting beyond the<br />

surface.<br />

ENDOSOME. A rather confused pattern <strong>of</strong> vague tracts, scattered spicules and a largely<br />

unispicular spongin-bound network. In the Port Royal specimens, sigmas are abundant<br />

throughout the sponge. Much <strong>of</strong> the interior <strong>of</strong> the cay sponges is occupied by algae.<br />

Sigmas are exceedingly rare, even in algae-free portions.<br />

SPICULES, (a) Oxeas, slightly curved, fusiform to wedge-shaped, 117-<strong>20</strong>0 X 5-9 p. Port<br />

Royal specimens, 128-<strong>20</strong>0 x 5-9 /A- Cay specimens, 117-177 x 3-5 /*,. Thinner develop-


SYSTEMATICS 31<br />

mental forms are common. A few styles <strong>of</strong> similar size occur, (b) Sigmas, thin, <strong>of</strong>ten<br />

sharply curved in the middle, with the ends nearly in one plane, 13-28 JU,. Individual<br />

analyses, lengths in microns:<br />

Oxeas<br />

143-<strong>20</strong>0 (50)*<br />

164-199 (50)<br />

128-182 (50)<br />

140-182 (50)<br />

140-188 (50)<br />

137-188 (50)<br />

145-188 (50)<br />

Port Royal Specimens<br />

Cay Specimens<br />

Sigmas<br />

17-23 (25)*<br />

22-25 (25)<br />

<strong>20</strong>-26 (25)<br />

<strong>20</strong>-30 (25)<br />

<strong>20</strong>-28 (25)<br />

15-<strong>20</strong> (25)<br />

15-26 (25)<br />

117-177 (50) 12-<strong>20</strong> (<strong>20</strong>)<br />

123-158 (50) 13-<strong>20</strong> (25)<br />

117-170 (50) 15-<strong>20</strong> (25)<br />

DISCUSSION. The varied development <strong>of</strong> a tangential dermal skeleton in this species<br />

indicates the close relationship <strong>of</strong> Sigmadocia and Gellius. The cay specimens are provisionally<br />

considered as conspecific with those <strong>of</strong> Port Royal. The type specimen is a<br />

sponge collected at Port Royal. The two lots <strong>of</strong> material agree in color and architecture.<br />

Oscular projections are, at best, weakly developed in the cay sponges which, however,<br />

grew in an area subject to considerable wave action. The cay sponges have thinner oxeas.<br />

The presence <strong>of</strong> rare sigmas similar in shape to those <strong>of</strong> the inshore specimens provides<br />

the best evidence <strong>of</strong> conspecificity.<br />

The species is noteworthy for its blue color and sharply curved sigmas. In spicule<br />

size, it is similar to some <strong>of</strong> the sponges attributed to Gellius fibulatus (Schmidt, 1862).<br />

Schmidt's original description provides little information on his species. The sponge<br />

photographed by Topsent (1925a) has a coarsely porous surface quite unlike that <strong>of</strong> my<br />

specimens. The sponge reported from Israel by LeVi (1957) differs from mine in having<br />

curious sigmas with a series <strong>of</strong> bends. Babic's specimens (1922), recorded as Gelliodes<br />

fibulata (Schmidt), had well-developed spongin fibers.<br />

FAMILY CALLYSPONGIIDAE de Laubenfels, 1936a<br />

GENUS CALLYSPONGIA Duchassaing and Michelotti<br />

Callyspongia fallax Duchassaing and Michelotti<br />

(Plate IV, fig. 2)<br />

Callyspongia fallax Duchassaing and Michelotti, 1864, p. 57 [type: St. Thomas; Brit. Mus.<br />

(Nat. Hist.) reg. no. 28.11.12.5]; Burton, 1934, p. 539; de Laubenfels, 1949, p. 13.<br />

HABITAT. Several specimens were collected in 10-15 feet <strong>of</strong> water near Maiden Cay.<br />

One was growing on the side <strong>of</strong> a Verongia fistularis, while the others were on coral<br />

masses.<br />

SHAPE. The sponge consists <strong>of</strong> a single, solid cylindrical branch, or an anastomosing<br />

group <strong>of</strong> such branches. The specimens range in length from 7-21 cm. The branches vary<br />

in diameter and also in outline along their length. They are usually 0.5-1.5 cm in width.<br />

A single specimen widens at one end to enclose a short vertical channel which is open<br />

at both ends. The cavity is lined by a smooth skin resembling that <strong>of</strong> the rest <strong>of</strong> the<br />

surface. It is occupied by a small crustacean and is certainly not a cloaca.<br />

COLOR. In life, the sponges are a dull purple. In alcohol, they become grayish-brown<br />

with or without traces <strong>of</strong> the original purple color.<br />

CONSISTENCY. Compressible.<br />

SURFACE. Smooth to the touch. The skin is even over wide areas <strong>of</strong> the surface. In


32 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

places, particularly distally, it is lifted into thorn-like conules, 2-4 mm high. Adjacent<br />

conules may be several millimeters or several centimeters apart. The oscules are usually<br />

scattered along the branches but may be sub-linear. They are 2-3 mm in diameter. Several<br />

exhalant canals open into each.<br />

ECTOSOME. An intergrading double network <strong>of</strong> spiculiferous spongin fibers. The coarse<br />

fibers have a diameter <strong>of</strong> 15-60 ju and contain 1-6 rows <strong>of</strong> spicules. They form an irregularly<br />

polygonal network, with parallel fibers 100-300 /x apart. The coarse mesh grades<br />

into a network <strong>of</strong> slender unispicular fibers which are 3-8 ^ in diameter. The fine meshes<br />

are usually less than a spicule length in width.<br />

ENDOSOME. The skeleton consists <strong>of</strong> uni- to multispicular spongin fibers, which are<br />

15-115 [A, but most frequently 30-60 JJL in diameter. The spicules may form a continuous<br />

core or may be spaced along the fiber. The nodes <strong>of</strong> fiber intersection are multispicular<br />

expansions, 70-115 ^ in diameter. Near the surface endosomal fibers run into the dermal<br />

mesh. The endosomal interstices are 100-300 fx in diameter and frequently rounded in<br />

outline. They contain scattered spicules similar in size and form to those <strong>of</strong> the fibers.<br />

SPICULES. Oxeas, straight to very slightly curved, with hastate or eroded points, 58-<br />

130 X 3-7 /A. The erosion may extend as far back as 2 diameters from the tip. Some<br />

spicules have low irregular swellings in mid-shaft. Individual analyses, lengths in microns,<br />

50 spicules each: 58-123; 70-130; 70-130.<br />

DISTRIBUTION. Tropical Atlantic America: Bahamas—de Laubenfels, 1949, p. 13;<br />

St. Thomas—Duchassaing and Michelotti, 1864, p. 57.<br />

DISCUSSION. The Jamaican solid, cylindrical, lavender Callyspongia is similar in appearance<br />

and architecture to the Bahaman sponges attributed by de Laubenfels to<br />

Callyspongia fallax. The spicules <strong>of</strong> my specimens, however, while small, are certainly<br />

not minute. Although C. fallax is the type-species <strong>of</strong> a large genus, little information<br />

is available on its structure. Duchassaing and Michelotti (1864) merely recorded specimens<br />

which, in their opinion, were conspecific with the unrecognizable Spongia papillaris<br />

<strong>of</strong> earlier authors. They established the name C. fallax, claiming that the name C.<br />

papillaris was preoccupied. Burton (1934) selected Duchassaing and Michelotti's specimen<br />

as the type <strong>of</strong> Callyspongia fallax. The type has been reexamined by Burton and<br />

probably by de Laubenfels as well. To my knowledge, neither author has published a<br />

detailed description <strong>of</strong> the sponge. Burton characterized the genus Callyspongia by its<br />

double dermal network, but curiously proceeded to emphasize the lack <strong>of</strong> such differentiation<br />

in the type-species, C. fallax.<br />

Burton's taxonomic summary (1934) cannot be accepted on the basis <strong>of</strong> published<br />

information. He rejects body form and spicule complement as valid specific criteria in<br />

the genus but does not present detailed evidence in support <strong>of</strong> this viewpoint. Judging<br />

from de Laubenfels' 1949 discussion, the type <strong>of</strong> C. fallax is solid and cylindrical. Burton<br />

could find no toxas in it. He considers Callyspongia eschrichtii Duchassaing and Michelotti,<br />

and Patuloscula procumbens Carter as synonyms <strong>of</strong> C. fallax, although he found<br />

toxiform spicules in them. In addition, both supposed synonyms are tubular or partially<br />

tubular sponges. Whatever the status <strong>of</strong> Duchassaing and Michelotti's Tuba scrobiculata,<br />

the species was established by Lamarck, whose material has been re-examined by Topsent,<br />

(1932). Callyspongia scrobiculata (Lamarck), a synonym <strong>of</strong> C. plicifera (Lamarck), can be<br />

distinguished from C. fallax by its tubular form, internal architecture, and strongylote<br />

spicules. Ceraochalina vanderhorsti Arndt, 1927, also included in the synonymy, is again<br />

basically tubular. The Jamaican sponges are clearly not conspecific with Arndt's species,<br />

from which they differ in having larger oxeas, a solid form and an abundance <strong>of</strong> spicules<br />

in the dermal network.<br />

Callyspongia vaginalis (Lamarck) de Laubenfels<br />

Spongia vaginalis Lamarck, 1813, p. 436 [type: American seas; Mus. nat. Hist. nat. Paris.]<br />

Tuba vaginalis, Duchassaing and Michelotti, 1864, p. 52.<br />

Callyspongia vaginalis, de Laubenfels, 1936a, p. 56; idem, 1949, p. 14; idem, 1950a, p. 56;<br />

idem, 1953a, p. 523; Wells et al., 1960, p. 210; Little, 1963, p. 41.<br />

[non] Callyspongia vaginalis, de Laubenfels, 1939, p. 2.


SYSTEMATICS 33<br />

Tuba sororia Duchassaing and Michelotti, 1864, p. 46 (fide de Laubenfels, 1936a); Carter,<br />

1884, p. <strong>20</strong>4.<br />

Spinosella sororia, Vosmaer, 1886, p. 342; Dendy, 1887d, p. 503; idem, 1890, p. 360;<br />

Wilson, 1902a, p. 394; Verrill, 1907, p. 334; Arndt, 1927, p. 155; Topsent, 1931, p. 75.<br />

[non] Spinosella sororia, Row, 1911, p. 326.<br />

HABITAT. Common on corals in 10-15 feet <strong>of</strong> water <strong>of</strong>fshore from the cays.<br />

SHAPE. The sponge may be a single, cylindrical tube or, more frequently, a group <strong>of</strong><br />

such tubes. The tubes have an even to jagged cloacal (oscular?) orifice, but never a papyraceous<br />

collar. The tubes are 1-3, occasionally 5 cm in diameter, and 5-15 cm in length.<br />

The width is rather uniform along the length <strong>of</strong> the tube. None <strong>of</strong> the specimens is a<br />

flaring vase. The walls are only a few millimeters in thickness. The cavities <strong>of</strong> adjacent<br />

tubes are <strong>of</strong>ten confluent at the base <strong>of</strong> the sponge.<br />

COLOR. The sponge is dull purple in life. In alcohol it retains much <strong>of</strong> its original<br />

color.<br />

CONSISTENCY. Compressible.<br />

SURFACE. The outer surface can vary between smooth and uneven along a tube. The<br />

elevations involve considerable areas <strong>of</strong> the surface. The inner wall has numerous openings,<br />

0.5-1 mm in diameter, and longitudinal ribs.<br />

ECTOSOME. A well-developed callyspongiid double network <strong>of</strong> spongin fibers. The<br />

coarse fibers are mostly <strong>20</strong>-30 /m in diameter and contain 2-4 rows <strong>of</strong> spicules. They outline<br />

irregularly polygonal spaces, 100-450 ^ in maximum width. The more symmetrical, finer<br />

network encloses apertures only a spicule or two in width. The fibers are 5-15 jn in diameter.<br />

They have a unispicular, usually spaced, core <strong>of</strong> oxeas. In the ectosome, the spicules<br />

are confined to the fibers. The cloacal surface has a coarse fiber network, which is condensed<br />

at the ribs into anastomosing, interwoven columns.<br />

ENDOSOME. The skeleton is formed by a reticulation <strong>of</strong> spongin fibers 15-75 ^ (mostly<br />

30-45 ju,) in diameter. All fibers have abundant spongin and most have a continuous core<br />

<strong>of</strong> several rows <strong>of</strong> spicules. Even large fibers may have only single, separated spicules<br />

over part <strong>of</strong> their length. The meshes are <strong>of</strong>ten nearly rectangular, with parallel fibers<br />

70-300 /x apart. A considerable number <strong>of</strong> spicules, mostly thin, are scattered in the flesh.<br />

The dermal network is supported by ascending fibers and <strong>of</strong>ten overlies subdermal<br />

cavities.<br />

SPICULES. Oxeas, 76-131 X 2-5 p. The spicules may be elongate and slightly curved, or<br />

short, straight and conspicuously hastate. The points are sometimes eroded. Individual<br />

analyses, lengths in microns, 50 spicules each: 83-123; 80-131; 76-123.<br />

DISTRIBUTION. Tropical Atlantic America: American Seas—Lamarck, 1813, p. 436 (as<br />

Spongia vaginalis); North Carolina—Wells et al., 1960, p. 210; Florida—Carter, 1884, p.<br />

<strong>20</strong>4 (as Tuba sororia); Dry Tortugas, Florida—de Laubenfels, 1936a, p. 56; West Coast,<br />

Florida—de Laubenfels, 1953a, p. 523; West Coast, Florida—Little, 1963, p. 41; Bermuda—<br />

Verrill, 1907, p. 334 (as Spinosella sororia) and de Laubenfels, 1950a, p. 56; Bahamas—<br />

Dendy, 1887d, p. 503; 1890, p. 360 (as Spinosella sororia), and de Laubenfels, 1949, p. 14;<br />

Puerto Rico—Wilson, 1902a, p. 324 (as Spinosella sororia); Lesser Antilles: Guadalupe, St.<br />

Thomas, St. Croix, Viecques, Tortola—Duchassaing and Michelotti, 1864, p. 52 (as Tuba<br />

vaginalis); St. Thomas, Viecques—Duchassaing and Michelotti, 1864, p. 46 (as Tuba<br />

sororia); Curasao—Arndt, 1927, p. 155 (as Spinosella sororia). Tropical Pacific America:<br />

Galapagos Is.—de Laubenfels, 1939, p. 2 (?).<br />

DISCUSSION. Lamarck's specimen has been restudied by both Topsent (1932) and de<br />

Laubenfels (1936a). The spicules <strong>of</strong> Jamaican specimens are rather robust for the species.<br />

Published data indicate that there is considerable variation in the size and abundance<br />

<strong>of</strong> the spicules. The fiber size <strong>of</strong> the Jamaican specimens agrees closely with that given<br />

in previous descriptions.<br />

The records <strong>of</strong> Duchassaing and Michelotti, Carter, and Verrill require verification.<br />

The two non-West Indian records are both very dubious. De Laubenfels' identification<br />

<strong>of</strong> the Galapagos Callyspongia was only tentative, since the specimen "does not agree<br />

well with ordinary examples <strong>of</strong> vaginalis." (1939, p. 2) Unfortunately, no other data were


34 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

provided. The Red Sea sponge studied by Row certainly differs from typical specimens<br />

<strong>of</strong> the West Indian C. vaginalis in interior structure. It had strong multispicular primary<br />

fibers, containing 10-<strong>20</strong> rows <strong>of</strong> spicules, and delicate, unispicular secondary fibers.<br />

De Laubenfels (1950a, p. 57) placed Siphonochalina stolonifera Whitfield, 1901, <strong>of</strong><br />

Bermuda into synonymy with C. vaginalis. Whitfield, however, described the spicules <strong>of</strong><br />

his sponge as cylindrical, which may mean strongylote. Schmidt's Siphonchalina papyracea<br />

(1870), <strong>of</strong>ten considered as a synonym <strong>of</strong> C. vaginalis, was described as having strongylote<br />

spicules ("umspitzigen nadeln"). Since a strongylote vase-shaped species, Callyspongia<br />

plicifera, occurs in the West Indies, the two cannot safely be taken as synonyms <strong>of</strong><br />

C. vaginalis.<br />

Callyspongia plicifera (Lamarck) de Laubenfels<br />

(Plate IV, fig. 1)<br />

Spongia plicifera Lamarck, 1813, p. 435 [type: American seas (?); Mus. nat. Hist. nat.<br />

Paris.]<br />

Tuba plicifera, Duchassaing and Michelotti, 1864, p. 53; Schmidt, 1870, p. 29; Carter,<br />

1882, p. 365.<br />

Spinosella plicifera, Dendy, 1887d, p. 506; idem, 1890, p. 365; Topsent, 1932, p. 73.<br />

Callyspongia plicifera, de Laubenfels, 1950a, p. 61.<br />

[non] Patuloscula plicifera, de Laubenfels, 1936a, p. 57 (fide de Laubenfels, 1950a, p. 61.)<br />

Spongia scrobiculata Lamarck, 1813, p. 436.<br />

Tuba scrobiculata, Duchassaing and Michelotti, 1864, p. 53.<br />

Spinosella scrobiculata, Topsent, 1932, p. 74.<br />

Spinosella maxima Dendy, 1887d, p. 506; idem, 1890, p. 365.<br />

HABITAT. Growing on coral masses in 10-15 feet <strong>of</strong> water <strong>of</strong>fshore from the cays.<br />

SHAPE. The sponge may be a single tube or a cluster <strong>of</strong> tubes. The individual tubes<br />

may be cylindrical or, more' frequently, vase-shaped. Small hollow, open buds are <strong>of</strong>ten<br />

present on the outside, and in one exceptional case, on the inside <strong>of</strong> the tube. The<br />

sponges are typically 10-30 cm in height. The cloacal (oscular?) orifice seldom has a<br />

diameter <strong>of</strong> less than 5 cm. In vase-shaped examples, the maximum diameter may exceed<br />

the length <strong>of</strong> the sponge. The walls are only a few millimeters in thickness.<br />

COLOR. A dull purple in life. In alcohol, the sponge may be a purplish or a dull<br />

yellow, or a mixture <strong>of</strong> the two colors. Dried specimens are beige.<br />

CONSISTENCY. Compressible.<br />

SURFACE. The surface becomes even toward the distal end. The terminus has a thin<br />

margin, in which a fiber reticulation is visible to the unaided eye. In some specimens,<br />

ribs composed <strong>of</strong> longitudinal fibers project beyond the general level <strong>of</strong> the cloacal<br />

terminus. The margin may be only a few millimeters or as much as several centimeters in<br />

length. The remainder <strong>of</strong> the surface may vary between uneven and strongly uneven on<br />

a single specimen. Many specimens are covered with broad blunt elevations which <strong>of</strong>ten<br />

grade into longitudinal or horizontal ridges. The elevations are several millimeters in<br />

height. The inner surface is even and pierced by numerous small oscules.<br />

ECTOSOME. An intergrading double network <strong>of</strong> spongin fibers which are mostly unispicular.<br />

The finer fibers are 15-30 ^ in diameter. They enclose meshes which are frequently<br />

150-300 fx in maximum width. Coarse fibers are typically 40-150 ^ in diameter,<br />

with parallel fibers <strong>of</strong>ten more than 0.5 mm apart. The longitudinal fibers <strong>of</strong> the top<br />

margin and fibrous projections are stout (70-225 fi in diameter) and frequently multispicular.<br />

In the latter case, the spicules form an axial core. The primaries are joined by<br />

less robust connectives and also by thin fibers similar to those <strong>of</strong> the endosome.<br />

ENDOSOME. The main skeleton is a wide-meshed, rounded to polygonal network <strong>of</strong><br />

spongin fibers. The fibers are 40-90 ^ or occasionally as much as 300 ^ in diameter. They<br />

may be unispicular, or have a multispicular axial core. The spicules may be spaced, or they<br />

may form a continuous line. A very irregular finer mesh joins the main skeleton and is<br />

<strong>of</strong>ten interwoven with it. Its fibers are 15-30 JJL in diameter and contain spaced single<br />

spicules.


SYSTEMATICS 35<br />

SPICULES. Strongyles, truncate, straight to somewhat curved or bent, 53-100 X 1-2 JJL.<br />

Individual analyses, lengths in microns, 50 spicules each: 67-97; 63-100; 68-86; 53-91.<br />

DISTRIBUTION. Tropical Atlantic America: American Seas—Lamarck, 1813, p. 435 (as<br />

Spongia plicifera); Florida—Schmidt, 1870, p. 29 (as Tuba plicifera); Bahamas—Dendy,<br />

1887d, p. 506; 1890, p. 365 (as Spinosella plicifera), Dendy, 1887d, p. 506; 1890, p. 365<br />

(as Spinosella maxima); Antilles—Duchassaing and Michelotti, 1864, p. 53 (as Tuba<br />

plicifera), Schmidt, 1870, p. 29 (as Tuba plicifera); Jamaica—Dendy, 1887d, p. 506, 1890,<br />

p. 365 (as Spinosella maxima); Lesser Antilles, St. Thomas, Tortole, Guadeloupe—<br />

Duchassaing and Michelotti, 1864, p. 53 (as Tuba scrobiculata), Grenada—Carter, 1882,<br />

p. 365 (as Tuba plicifera),<br />

DISCUSSION. The Jamaican specimens are similar in architecture and spiculation to<br />

Spongia scrobiculata Lamarck, as redescribed by Topsent. Burton (1934, p. 539) attributed<br />

that species to Duchassaing and Michelotti and considered it as a synonym <strong>of</strong><br />

Callyspongia fallax. Their specimen <strong>of</strong> scrobiculata was described as tubular, and thus is<br />

unlikely to be conspecific with C. fallax. Spongia scrobiculata and S. plicifera Lamarck<br />

have been treated as closely related species by Lamarck, Duchassaing and Michelotti, and<br />

Topsent. The latter species is characterized by well-developed, largely horizontal folds<br />

and grooves. According to Topsent (1932) spicules are absent in the type. S. scrobiculata<br />

also has surface convolutions which outline deep fossae in Lamarck's specimens. Topsent<br />

found thin strongylote spicules in the fibers <strong>of</strong> the fossa-bearing specimens. He also<br />

found strongyles in a specimen (photographed by Topsent, 1932, plate 2, figure 4) which<br />

Lamarck had assigned to Spongia plicifera. The specimen has low, irregular to horizontal<br />

folds, making it very similar to my material. Topsent considered the sponge to be an<br />

example <strong>of</strong> scrobiculata, using the presence or absence <strong>of</strong> spicules as a distinguishing<br />

characteristic.<br />

A collection <strong>of</strong> Cuban specimens (CU-6, CU-6A-E) on loan to the <strong>Yale</strong> <strong>Peabody</strong> <strong>Museum</strong><br />

from the U. S. National <strong>Museum</strong> strongly suggests that the two species are actually<br />

conspecific. The dried Cuban sponges are a pale beige in color. They have a thin distal<br />

margin as in the Jamaican sponges. On the basis <strong>of</strong> surface folds and depressions all <strong>of</strong><br />

the Cuban specimens must be assigned to Callyspongia plicifera. CU-6C, for example, is<br />

identical in appearance with the specimens <strong>of</strong> plicifera figured by Dendy and by Duchassaing<br />

and Michelotti. The surface lamellae are well-developed in all and are almost entirely<br />

horizontal in CU-6C. The folds become irregular and meandriform over part or<br />

most <strong>of</strong> the surface in CU-6B and CU-6. Strongly convoluted Jamaican specimens certainly<br />

approach that condition.<br />

Strongylote spicules <strong>of</strong> 2 p, diameter are present in all <strong>of</strong> the Cuban examples. They<br />

are similar in shape and size to those <strong>of</strong> the Jamaican sponges. For example, a sample <strong>of</strong><br />

50 spicules in CU-6 varied in length between 68 and 94 p. Topsent, Schmidt, and Dendy<br />

were unable to find spicules in their examples <strong>of</strong> C. plicifera. Carter, however, reported<br />

very thin acerate spicules, 72 /x in length, for his Grenada specimen. The Cuban specimens<br />

have a skeletal pattern quite comparable to that found in the Jamaican sponges.<br />

The surface is made up <strong>of</strong> a coarse double network. The main skeleton <strong>of</strong> the interior is<br />

constructed <strong>of</strong> stout spongin fibers (35-150 ^ in diameter). They may contain only single<br />

spicules or a multispicular axial core. Several <strong>of</strong> the specimens (CU-6A, CU-6E) have fewer<br />

spicules than the Jamaican sponges. The main skeleton is joined at intervals by a very<br />

irregular network <strong>of</strong> fine fibers. The latter are only about 15 ^ in diameter and contain<br />

very few spicules. Dendy (1890) reported a similar network in his specimens <strong>of</strong> plicifera.<br />

In summary, the Jamaican and Cuban specimens differ only in the extent to which the<br />

surface is thrown into horizontal convolutions.<br />

Spinosella maxima Dendy, 1887d, is very probably conspecific with Callyspongia<br />

plicifera. The outer surface is uneven except for a thin distal margin. The interior<br />

skeleton had the same pattern as in plicifera, including an irregular network <strong>of</strong> delicate<br />

fibers. Spicules were said to be absent from the Bahaman examples, but thin vestigial<br />

oxeas were reported for a Jamaican specimen.<br />

Siphonochalina papyracea Schmidt, 1870, <strong>of</strong>ten considered as a synonym <strong>of</strong> Callyspongia<br />

vaginalis, is more likely to prove to be conspecific with C. plicifera. The external


36 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

appearance is certainly similar, and the spicules, described as "umspitzigen nadeln," are<br />

apparently strongylote.<br />

Callyspongia pallida n. sp.<br />

Fig. 6. —Spicules <strong>of</strong> Callyspongia pallida. Oxeas, style, strongyle. YPM 5038. Holotype.<br />

HOLOTYPE. YPM 5038. Seawall <strong>of</strong> Police Post. November 30, 1960.<br />

HABITAT. The specimen was collected on the seawall <strong>of</strong> the Police Post at Port Royal<br />

in several feet <strong>of</strong> water.<br />

SHAPE. The sponge is a horizontally spreading complex <strong>of</strong> lobes. The lobes are 1.5-2<br />

cm in height and have apical oscules. Connecting stems are about 1 cm long and 0.5-1 cm<br />

in diameter.<br />

COLOR. The sponge was dull yellow in life and is a pale beige in alcohol.<br />

CONSISTENCY. Compressible, s<strong>of</strong>t, elastic.<br />

SURFACE. Even, smooth to the touch.<br />

ECTOSOME. The dermal layer can be detached only with difficulty. It is a callyspongiid<br />

double network <strong>of</strong> spongin fibers. The two networks gradually intergrade with respect to<br />

fiber diameter. The small fibers are 7-30 LL in diameter. They are mostly unispicular,<br />

with the spicules in a continuous line, or less frequently, somewhat spaced. The meshes<br />

are 60-2<strong>20</strong> LL in width. They are <strong>of</strong>ten nearly square or rectangular, with rounded corners.<br />

The large fibers are 30-45 ^ in diameter and usually have several rows <strong>of</strong> spicules in a<br />

continuous axial core. The meshes are triangular to polygonal. Parallel fibers may be<br />

<strong>20</strong>0-580 LL apart. The dermal pores are 35-45 LL in diameter. A few interstitial spicules<br />

are present.<br />

ENDOSOME. The interior fibers are <strong>20</strong>-75 LL in diameter. Thinner fibers are mostly<br />

unispicular. Thicker fibers have a continuous axial core <strong>of</strong> 3-6 rows. The square to rectangular<br />

meshes are mostly 300-600 LL in maximum width. A few interstitial spicules<br />

occur in the flesh. A number <strong>of</strong> dense, nearly spherical embryos, 160-217 LL in diameter,<br />

are present in the specimen which was collected in November.<br />

SPICULES. Mostly diactinal, straight to very slightly curved, 48-102 X 1-5 LL.* (125<br />

spicules.) Most <strong>of</strong> the spicules are centrotylote, with a central irregularly swollen area<br />

about 7 LL long, and 3-5 LL in diameter. The ends are most frequently narrow but blunt.<br />

Others are acutely pointed, irregular in outline, or round. In the last case, the spicules<br />

are styles or strongyles. The modified spicules may be centrotylote. The spicules <strong>of</strong> the<br />

interior fibers are <strong>of</strong>ten very thin, with the shaft (aside from any central swelling) only<br />

1-2 LL in diameter.


SYSTEMATICS 37<br />

DISCUSSION. The most closely related species is Callyspongia aspinosa Levi, 1959, <strong>of</strong><br />

West Africa. They are similar in appearance, color, and consistency. Levi's sponge is<br />

more tubular in shape. The spicules <strong>of</strong> the West African sponge are substrongyles which<br />

may reach 125 p in length. The dimensions <strong>of</strong> the interior skeleton are somewhat<br />

different.<br />

ORDER POECILGSCLERIDA Topsent 1928<br />

FAMILY TEDANIIDAE Ridley and Dendy, sensu de Laubenfels, 1936a<br />

GENUS TEDANIA Gray<br />

Tedania ignis (Duchassaing and Michelotti) Verrill<br />

(Plate V, fig. 4)<br />

Thalysias ignis Duchassaing and Michelotti, 1864, p. 83 [type: St. Thomas; Brit. Mus.<br />

(Nat. Hist.) reg. no. 28.11.12.43.]<br />

Tedania ignis, Verrill, 1907, p. 339; de Laubenfels, 1936a, p. 89; idem, 1936b, p. 459;<br />

idem, 1949, p. 16; idem, 1950a, p. 69; idem, 1950b, p. 21 (?); idem, 1951a, p. 260 (?);<br />

idem, 1956, p. 3; Little, 1963, p. 48.<br />

Tedania ignis pacifica de Laubenfels, 1954a, p. 129 (?).<br />

Tedania digitata, Schmidt (not Schmidt, 1862), 1870, p. 43 [partim]; Wilson, 1902a, p.<br />

395 (fide de Laubenfels, 1936a); Arndt, 1927, p. 146.<br />

Tedania digitata var. bermudensis Ridley and Dendy, 1887, p. 51.<br />

Reniera digitata, Carter, 1882, p. 287; idem, 1884, p. <strong>20</strong>5.<br />

Tedania brucei Wilson, 1894, p. 3<strong>20</strong> (fide de Laubenfels, 1936a.)<br />

Tedania nigrescens, Burton and Rao, 1932, p. 353 [partim] (not Schmidt, 1862.)<br />

Tedania anhelans, Burton, 1954, p. 229 (not Lieberkiihn, 1859.)<br />

HABITAT. Common on mangrove roots, pilings at Port Royal and on coral rubble<br />

near the cays.<br />

SHAPE. Thickly incrusting, and massive to lobate. Typical specimens are somewhat<br />

larger than fist-sized. Massive specimens have prominent volcano-like oscular projections.<br />

COLOR. The exterior is a very uniform flame red. The interior is a pale red-orange.<br />

In alcohol the sponges become beige.<br />

CONSISTENCY. Compressible. De Laubenfels <strong>of</strong>ten described the species as extremely<br />

s<strong>of</strong>t and delicate, but that is certainly not the case in the Jamaican specimens.<br />

SURFACE. The smooth dermal skin is made uneven by low meandering ridges and<br />

conules. The numerous oscules, <strong>of</strong>ten raised on projections, are several millimeters to<br />

1 cm in diameter.<br />

ECTOSOME. The skin contains thickly strewn tylotes and onychaetes. The tylotes <strong>of</strong>ten<br />

form broad but ill-defined tracts. The scattered ostia are 40-60 p in diameter.<br />

ENDOSOME. The skeleton includes styles grouped into spicule tracts 40-70 //, in diameter.<br />

The tracts grade into loose bundles and scattered spicules. Onychaetes are<br />

thickly strewn in the flesh. Large gemmules, 300-500 ^ in longest diameter, are present<br />

in the interior. They are dense aspiculous masses, bright red in life, and white in alcohol.<br />

All <strong>of</strong> the dozens <strong>of</strong> specimens examined in the field at all times <strong>of</strong> the year had numerous<br />

gemmules. Their development into larvae has been followed by Wilson (1894).<br />

SPICULES, (a) Dermal tylotes, straight, with narrow, elongate heads, 181-253 X 3-7 //,.<br />

The heads are terminally microspined. (b) Styles, smooth, straight to slightly curved, with<br />

narrowed heads and rather hastate points, 232-297 X 5-12 /x. (c) Onychaetes, microspined<br />

raphides, 47-264 X 1-3 p.. It is difficult to determine the lower range. Individual analyses,<br />

lengths in microns, 25 spicules each:<br />

Tylotes<br />

214-236<br />

199-234<br />

<strong>20</strong>5-234<br />

210-232<br />

210-253<br />

181-246<br />

Styles<br />

253-293<br />

234-297<br />

234-281<br />

232-270<br />

239-283<br />

232-283<br />

Raphides<br />

47-258<br />

159-253<br />

<strong>20</strong>3-264<br />

87-232<br />

137-253<br />

101-239


38 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

DISTRIBUTION. Tropical Atlantic America: Florida—Carter, 1884, p. <strong>20</strong>5 (as Reniera<br />

digitata); Dry Tortugas, Florida—de Laubenfels, 1936a, p. 89; West coast, Florida—<br />

Little, 1963, p. 48; Atlantic coast, Panama—de Laubenfels, 1936b, p. 459; Brazil—de<br />

Laubenfels, 1956, p. 3; Bermuda—Ridley and Dendy, 1887, p. 51 (as Tedania digitata<br />

bermudensis), Verrill, 1907, p. 339, de Laubenfels, 1950a, p. 69; Bahamas—Wilson, 1894,<br />

p. 3<strong>20</strong> (as Tedania brucei), de Laubenfels, 1949, p. 16; Puerto Rico—Wilson, 1902a, p.<br />

395 (as Tedania digitata), Lesser Antilles—St. Thomas—Duchassaing and Michelotti,<br />

1864, p. 83 (as Thalysias ignis), Schmidt, 1870, p. 43 (as Tedania digitata)) Antigua—<br />

Carter, 1882, p. 287 (as Reniera digitata); Curasao—Arndt, 1927, p. 146 (as Tedania<br />

digitata); Turneffe Is. (West Indies)—Burton, 1954, p. 229 (as Tedania anhelans); Central<br />

Pacific—de Laubenfels, 1954a, p. 129 (?); Hawaiian Is.—de Laubenfels, 1950b, p.<br />

21, 1951a, p. 260 (?).<br />

DISCUSSION. Specimens <strong>of</strong> Tedania with similar-sized spicules are widely distributed<br />

in warm seas. Burton and Rao (1932) concluded that all were conspecific with the<br />

Mediterranean sponge now known as Tedania anhelans (Lieberkiihn). De Laubenfels<br />

(1936a, 1950a, and 1950b) maintained that the West Indian population was specifically<br />

distinct from the European one, largely on the basis <strong>of</strong> differences in coloration and<br />

style thickness. A review <strong>of</strong> the literature casts doubt on the latter criterion. The uniformity<br />

<strong>of</strong> coloration in the West Indian examples <strong>of</strong> Tedania is most striking. For example,<br />

all <strong>of</strong> the many dozens <strong>of</strong> Tedania ignis seen in a variety <strong>of</strong> shallow water habitats<br />

over a two-year period in Jamaica had the same flame-red color. De Laubenfels noted<br />

only three variants in a population <strong>of</strong> "literally thousands" at the Dry Tortugas. (1936a,<br />

p. 90.) A study <strong>of</strong> eastern Caribbean faunas might provide information <strong>of</strong> value, since<br />

Schmidt recorded both black and red sponges from St. Thomas. By contrast, the Mediterranean<br />

and West African sponges are variously reported as green, orange, gray, brown<br />

and black. A branching growth form has been recorded for Tedania from Europe but<br />

has not been found in the West Indian sponges. Tedania ignis is said to irritate<br />

severely the skin <strong>of</strong> many people, although it did not affect me. Such irritation has not,<br />

to my knowledge, been associated with the Mediterranean sponge. Gemmules have been<br />

reported only for the West Indian sponge (cf. Wilson 1894, de Laubenfels 1950a) and<br />

may be a characteristic <strong>of</strong> T. ignis. The West Indian population, therefore, forms a<br />

rather homogeneous group, which is very probably not conspecific with the Mediterranean<br />

T. anhelans.<br />

Specimens <strong>of</strong> Tedania from the Indo-Pacific also present a taxonomic problem. De<br />

Laubenfels only doubtfully assigned Hawaiian and Central Pacific sponges to the West<br />

Indian species. His Pacific specimens were always incrusting. He concluded that they<br />

were at least a distinct subspecies. Australasian specimens <strong>of</strong> Tedania, recorded under<br />

a number <strong>of</strong> species names, are described as orange or red in color. Their relationship<br />

to the European and West Indian species remains to be clarified.<br />

GENUS LISSODENDORYX Topsent<br />

Lissodendoryx isodictyalis (Carter) Topsent<br />

Halichondria. isodictyalis Carter, 1882, p. 285 [type: Puerto Cabello, Venezuela, and<br />

Acapulco, Mexico; Brit. Mus. (Nat. Hist.) (?)]; idem, 1886, p. 52.<br />

Myxilla isodictyalis, Dendy, 1896, p. 30; Row, 1911, p. 343.<br />

Lissodendoryx isodictyalis, Topsent, 1897, p. 456; idem, 1925a, p. 701; idem, 1936, p. 18;<br />

Burton, 1948, p. 755; de Laubenfels, 1936a, p. 93; idem, 1947, p. 35; idem, 1950a, p.<br />

73; idem, 1956, p. 2, 3; Dickinson, 1945, p. <strong>20</strong>; L£vi, 1952, p. 48; Hopkins, 1956, p. 24;<br />

Hartman, 1958, p. 41; Wells et al., 1960, p. 212; Sara and Siribelli, 1960, p. 53; Little,<br />

1963, p. 48.<br />

Lissodendoryx isodictyalis paucispinosa Topsent, 1928, p. 240 (?).<br />

[non] Lissodendoryx isodictyalis jacksoniana Burton, 1936b, p. 141.<br />

Tedania leptoderma Topsent, 1889, p. 49 (fide Topsent, 1897.)<br />

Lissodendoryx leptoderma, Topsent, 1894, p. 35.<br />

Lissodendoryx similis Thiele, 1899, p. 18 (fide Topsent, 1925a); Burton and Rao, 1932,<br />

p. 331 (fide Topsent, 1936.)


SYSTEMATICS 39<br />

Lissodendoryx carolinensis Wilson, 1911, p. 11; George and Wilson, 1919, p. 150; Mc-<br />

Dougall, 1943, p. 331.<br />

HABITAT. The single specimen was collected at Drunkenman's Cay, at a turtle grass<br />

bed in three feet <strong>of</strong> water.<br />

SHAPE. The sponge is a low cushion-shaped mass 4.5 X 3 cm in area. It attains a maximum<br />

thickness <strong>of</strong> two cm.<br />

COLOR. In life, light purple. In alcohol the sponge is nearly cream.<br />

CONSISTENCY. Spongy, compressible.<br />

SURFACE. The dermis is a detachable skin. Its low, meandering convolutions are<br />

similar to those <strong>of</strong> Tedania ignis. A few oscules, 1-5 mm in diameter, are scattered over<br />

the surface.<br />

ECTOSOME. The dermal membrane contains tylotes and abundant microscleres. The<br />

tylotes are strewn singly and in loose bundles, 30-40 /m in diameter. Dermal pores pierce<br />

the skin singly and in groups <strong>of</strong> three to six. Typical ostial sizes are 50 X 35, 87 X 72,<br />

and 116 x 94 ^. Extensive subdermal cavities, 100-300 LL in depth, underlie the skin.<br />

The surface layer is supported by broad tufts <strong>of</strong> tylotes which begin as bundles below the<br />

subdermal spaces.<br />

ENDOSOME. The styles form a confused to subisodictyal pattern in the interior. Illdefined<br />

spicule tracts are present in places. A few tylotes are present in the floor <strong>of</strong> the<br />

subdermal spaces. Microscleres are extremely abundant. The chelas are <strong>of</strong>ten clustered<br />

into groups and lines. A few clumps <strong>of</strong> debris were noted in the flesh.<br />

SPICULES, (a) Dermal tylotes, straight, with swollen elongate ends, 188-223 X 3-5 LL<br />

(50 spicules), (b) Endosomal styles, usually with the shaft curved near the basal end,<br />

152-188 X 3-7 fi. The spicules are somewhat subtylostylote, with a slight neck constriction<br />

and narrow head. Their points are rather hastate, narrowing from within lV^-2<br />

diameters <strong>of</strong> the end (50 spicules), (c) Sigmas, 22-28 /* (30 spicules), (d) Tridentate (arcuate)<br />

isochelas, with a slightly curved shaft, 27-37 LL (30 spicules). The central teeth are<br />

each 10 [x long, in a spicule <strong>of</strong> 27 LL total length. The ends have narrow but definitely<br />

rounded apices.<br />

DISTRIBUTION. North American temperate Atlantic: New England—Hartman, 1958,<br />

p. 41.<br />

Tropical Atlantic America: North Carolina—Wilson, 1911, p. 11. (as Lissodendoryx<br />

carolinensis), George and Wilson, 1919, p. 150, (as Lissodendoryx carolinensis), Mc-<br />

Dougall, 1943, p. 331 (as Lissodendoryx carolinensis), de Laubenfels, 1947, p. 35, Wells<br />

et al., 1960, p. 212; South Carolina—Hopkins, 1956, p. 24; Dry Tortugas, Florida—de<br />

Laubenfels, 1936a, p. 93; West coast, Florida—Little, 1963, p. 48; Venezuela—Carter,<br />

1882, p. 285 (as Halichondria isodictyalis); Brazil—de Laubenfels, 1956, p. 2, 3; Bermuda—de<br />

Laubenfels, 1950a, p. 73, Hartman, 1958, p. 41; Guadeloupe—Topsent, 1889,<br />

p. 49 (as Tedania leptoderma).<br />

Mediterranean-Atlantic: Monaco—Topsent, 1925a, p. 701, Topsent, 1936, p. 18;<br />

Naples, Italy—Topsent, 1925a, p. 701, Sara and Siribelli, 1960, p. 53; Venice, Italy—<br />

Topsent, 1925a, p. 701.<br />

Tropical West Africa: Cape Verde Is.—Topsent, 1928, p. 240 (?); Senegal, LeVi, 1952,<br />

p. 48; Congo—Burton, 1948, p. 755.<br />

Indo-Pacific-Red Sea—Row, 1911, p. 343 (as Myxilla isodictyalis); Mergui Archipelago,<br />

<strong>of</strong>f Burma—Burton and Rao, 1932, p. 311 (as Lissodendoryx similis); East Indies—<br />

Topsent, 1897, p. 456; Thiele, 1899, p. 18 (as Lissodendoryx similis); Australia—Carter,<br />

1886, p. 52 (as Halichondria isodictyalis); Dendy, 1896, p. 30 (as Myxilla isodictyalis);<br />

Whitelegge, 1902, p. 25 (as Myxilla isodictyalis);<br />

Tropical Pacific America: Acapulco, Mexico—Carter, 1882, p. 285 (as Halichondria<br />

isodictyalis); Gulf <strong>of</strong> California—Dickinson, 1945, p. <strong>20</strong>.<br />

DISCUSSION. A variety <strong>of</strong> colors has been reported for this species, but the present<br />

specimen is the first purple example. The unpleasant Ircinia-like odor reported by<br />

de Laubenfels (1950a), and apparently noticed by Hopkins, who called it the garlic sponge,<br />

was noted in the Jamaican sponge.


40 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

In spiculation, my specimen agrees well with Topsent's Guadeloupe sponge, and,<br />

except for sigma size, with Carter's type. The various specimens attributed to the species<br />

exhibit a wide range <strong>of</strong> spicule sizes. The reported variations cannot be correlated with<br />

distribution. For example, wide extremes <strong>of</strong> microsclere size have been reported for both<br />

West Indian and Mediterranean specimens. Sponges from the East Coast <strong>of</strong> the United<br />

States do form a rather homogeneous group, characterized by small chelas and large<br />

sigmas (as compared with the known ranges for the species). They also have peculiar,<br />

contractile, pore-bearing dermal papillae. Hartman (1958) has referred to them as the<br />

carolinensis type, and they could be considered as a separate subspecies.<br />

Lissodendoryx similis Thiele has spicules similar in size to those <strong>of</strong> L. isodictyalis.<br />

Thiele emphasized the presence <strong>of</strong> chelas with round rather than pointed ends in his<br />

species definition. Topsent (1925a, 1936) concluded that chela shape was a variable characteristic<br />

within L. isodictyalis.<br />

Lissodendoryx jacksoniana (Lendenfeld, 1888), as redescribed by Hallmann (1914),<br />

differs sufficiently from L. isodictyalis to warrant its retention as a separate species. Its<br />

endosomal skeleton includes small strongyles in addition to the usual styles. Burton<br />

(1936b) has reported it from South Africa as L. isodictyalis jacksoniana.<br />

The poorly known Lissodendoryx isodictyalis paucispinosa Topsent is similar in<br />

spicule sizes to Carter's species. It approaches the myxillid condition with varying degrees<br />

<strong>of</strong> spination on the base and even the shaft <strong>of</strong> the styles.<br />

GENUS ACANTHACARNUS Levi<br />

Acanthacarnus souriei Levi<br />

Acanthacarnus souriei Levi, 1952, p. 54 [type: Senegal; repository not specified]; idem,<br />

1959, p. 132; Vacelet, 1961, p. 42.<br />

Acanthacarnus levii Vacelet, 1960, p. 267 (fide Vacelet, 1961.)<br />

HABITAT. The single example was collected at Drunkenman's Cay.<br />

SHAPE. A very thin film, in crevices <strong>of</strong> a coral rock.<br />

COLOR. Bright red-orange.<br />

CONSISTENCY. TOO thin to determine.<br />

SURFACE. Smooth to the touch, even. No oscules are apparent.<br />

ECTOSOME. The dermal tylotes are thickly strewn at the surface. Toxas and chelas are<br />

abundant.<br />

ENDOSOME. The shape <strong>of</strong> the specimen made sectioning difficult. A few bundles <strong>of</strong><br />

styles, about 30 fx in diameter, could be seen. They are obliquely echinated by cladotylotes<br />

and acanthostyles. The cladome end <strong>of</strong> the cladotylotes and the apices <strong>of</strong> the<br />

acanthostyles are directed outward from the spicule bundles.<br />

SPICULES, (a) Dermal tylotes, straight, heads usually microspined, 119-357 X 3-7 /x (50<br />

spicules), (b) Endosomal styles, <strong>of</strong>ten subtylostylote, slightly curved, 240-381 X 3-5 xi<br />

(30 spicules). Their bases are slightly swollen and roughened, or irregularly shaped.<br />

(c) Cladotylotes (rose-stem spicules), straight to slightly curved, 70-<strong>20</strong>5 x 3-5 /x (60 spicules).<br />

One end has four well-developed recurved clads. The other end may have short<br />

irregularly oriented clads or lumpy projections. The stem bears numerous thorn-like<br />

spines, which usually point away from the cladome. (d) Echinating acanthostyles, thin,<br />

straight, finely and entirely spined, 68-104 x 3-5 ^ (30 spicules), (e) Palmate isochelas,<br />

17-21 X 2 /a (50 spicules), (f) Toxas, varied in shape, 50 X 2 to 330 X less than 2 /x.<br />

The more elongate toxas are thin, with a short, shallow central arch. The smaller ones<br />

are thick and strongly arched.<br />

DISTRIBUTION. Mediterranean-Atlantic: Corsica—Vacelet, 1960, p. 267 (as Acanthacarnus<br />

levii); Vacelet, 1961, p. 42.<br />

Tropical West Africa: Senegal—Levi, 1952, p. 54; Gulf <strong>of</strong> Guinea—Levi, 1959, p. 132.<br />

DISCUSSION. The Jamaican specimen is the first West Indian record <strong>of</strong> the species. My<br />

specimen agrees closely with the West African examples.


n i<br />

SYSTEMATICS 41<br />

FAMILY MICROCIONIDAE Hentschel, sensu de Laubenfels, 1936a<br />

GENUS MICROCIONA Bowerbank<br />

Microciona microchela n. sp.<br />

10//<br />

c— -<br />

A 3H I 1<br />

100//<br />

Fig. 7. —Spicules <strong>of</strong> Microciona microchela. A. Tylostyle. B. Head <strong>of</strong> subtylostyle enlarged to<br />

show spination. C. Acanthostyle. D. Toxa <strong>of</strong> two sizes. E. Palmate isochelas <strong>of</strong> two<br />

sizes. C, D: scale I. B, E: scale II. A: scale III. YPM 5040. Holotype.<br />

HOLOTYPE. YPM 5040. Rasta's Wreck, May 24, 1961.<br />

HABITAT. Common on shells and pilings at Port Royal.<br />

SHAPE. Most specimens are incrustations about one mm in thickness. They <strong>of</strong>ten cover<br />

several square feet <strong>of</strong> the substratum. One specimen is an anastomosing complex <strong>of</strong> flattened,<br />

repent branches attached to the substratum at several places. Its branches are 1-5<br />

mm in maximum diameter. It is identical to the incrusting specimens in spiculation.<br />

COLOR. In life the sponges have a pale pink surface and a pinkish-orange interior. In<br />

alcohol they first become bluish-gray and finally beige to brown.<br />

CONSISTENCY. Tough, very slightly compressible.<br />

SURFACE. Smooth to the touch; even or partly uneven. River-like systems <strong>of</strong> narrow,<br />

efferent subdermal canals lead to small oscules. Neither canals nor events are evident in<br />

preserved specimens.<br />

ECTOSOME. The dermal palisade is difficult to detach. It consists <strong>of</strong> radiating tufts <strong>of</strong><br />

short, thin tylostyles. The spicules project slightly beyond the dermal membrane. The<br />

latter contains an abundance <strong>of</strong> minute isochelas, as well as smaller numbers <strong>of</strong> the other<br />

microsclere types. The dermal pores, clustered in groups <strong>of</strong> 4-8, are <strong>20</strong>-45 //, in diameter.<br />

ENDOSOME. Small acanthostyles and long, robust subtylostyles are embeddedd in a thin<br />

basal layer <strong>of</strong> spongin. Short spongin columns, with a diameter <strong>of</strong> 15-75 p, ascend from<br />

the basal plate in the incrusting specimens. The interior <strong>of</strong> the branching sponge is supported<br />

by an irregular fibroreticulation. The columns and fibers contain one to many<br />

rows <strong>of</strong> subtylostyles. They are echinated by acanthostyles and long, robust subtylostyles.<br />

The flesh contains all categories <strong>of</strong> microscleres and numerous long, thin subtylostyles.<br />

SPICULES, (a) Subtylostyles, straight, gradually pointed, 106-410 x 2-10 /x. Many spicules<br />

<strong>of</strong> all lengths have finely microspined or roughened heads. The dermal spicules are<br />

2-5 juL in diameter. The long, robust echinators are 5-10 fi in diameter. A few <strong>of</strong> the


42 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

robust spicules are styles, (b) Acanthostyles, straight, with sharp points, 50-100 ^ in length.<br />

The heads are 5-8 /x in diameter. Conical spines, 1-3 LL in height, are irregularly strewn<br />

over the head and basal 2/3-4/5 <strong>of</strong> the shaft, (c) Large palmate isochelas, 10-18 LL in<br />

length, with a maximum head diameter <strong>of</strong> 3 LI. (d) Small palmate isochelas, 4-7 LL in<br />

length, with a maximum head diameter <strong>of</strong> 2 ju,. (e) Large toxas, 48-413 X 1-2 fx. The<br />

longer spicules have a short, shallow central arch and straight sides. A very few are<br />

actually raphides. The short spicules have a deeper arch and slightly recurved ends.<br />

Toxodragmata are present, (f) Small toxas, 5-13 X 1-2 /A, with straight sides. The spicules<br />

approach an inverted, flattened V in shape. Individual analyses, lengths in microns:<br />

Subtylostyles<br />

110-386 (70)<br />

106-410 (70)<br />

117-351 (25)<br />

104-340 (50)*<br />

Small isochelas<br />

4-5 (25)<br />

5-7 (25)<br />

4-7 (25)<br />

5-7 (25)*<br />

Acanthostyles<br />

50-100 (50)<br />

53-73 (50)<br />

50-78 (25)<br />

50-80 (25)*<br />

Large toxas<br />

present<br />

48-270 (15)<br />

136-413 (13)<br />

48-3<strong>20</strong> (15)*<br />

Large isochelas<br />

10-13 (25)<br />

10-17 (25)<br />

10-16 (25)<br />

13-18 (25)*<br />

Small toxas<br />

present<br />

7-10 (13)<br />

present<br />

5-13 (25)*<br />

DISCUSSION. The Jamaican sponge is characterized by the presence <strong>of</strong> two sizes <strong>of</strong> both<br />

toxas and chelas. Microciona parthena de Laubenfels <strong>of</strong> California (cf. de Laubenfels,<br />

1932b) has small toxas which are similar in shape to those <strong>of</strong> the present specimens.<br />

The Californian species is otherwise quite different.<br />

Only a few species <strong>of</strong> Microciona have minute isochelas in addition to the typical<br />

isochelas <strong>of</strong> the genus. Clathria favosa Whitelegge, 1906, <strong>of</strong> Australia has such spicules,<br />

but lacks minute toxas. It is a branching sponge with a honeycombed surface. Clathria<br />

copiosa Topsent, var. curacaoensis Arndt, 1927, also has minute isochelas. It is certainly<br />

not conspecific with Topsent's material. Arndt's Curasao sponge differs from the Jamaican<br />

in having styles rather than subtylostyles. Its acanthostyles are entirely spined and smaller<br />

in size. Small toxas are not present. Microciona maunaloa de Laubenfels (1951a) <strong>of</strong><br />

Hawaii is similar in color, shape and structure to the Jamaican species. The long toxas<br />

are <strong>of</strong> similar form. M. maunaloa differs in lacking small toxas. Its subtylostyles are<br />

devoid <strong>of</strong> spines, and its acanthostyles are entirely spined. The differences, combined<br />

with the geographical gap, suggest that the Jamaican sponge is not conspecific with M.<br />

maunaloa. They are certainly closely related species.<br />

Microciona rarispinosa n. sp.<br />

(Text-fig. 8)<br />

HOLOTYPE. YPM 5041. Rasta's Wreck, June 24, 1961.<br />

HABITAT. Incrusting on pilings and mussel shells at Port Royal.<br />

SHAPE. All <strong>of</strong> the several dozen observed specimens were very thin films. Most were<br />

less than 0.5 mm in thickness.<br />

COLOR. The living sponges are bright red. Preserved specimens are drab to gray.<br />

CONSISTENCY. S<strong>of</strong>t.<br />

SURFACE. Even, microhispid, punctiform. No oscules were visible.<br />

ECTOSOME. No specialization. Many <strong>of</strong> the tylostyles project beyond the protoplasmic<br />

surface.<br />

ENDOSOME. The spongin skeleton consists <strong>of</strong> a basal plate and short ascending fibers<br />

which are 30-45 fx in diameter. The plate and fibers lack coring spicules but are echinated<br />

by stout subtylostyles. The echinators are basally implanted in spongin, <strong>of</strong>ten for i/s <strong>of</strong><br />

their length. The shorter echinators usually have roughened bases. The flesh contains<br />

both thick and thin subtylostyles <strong>of</strong> all sizes along with numerous microscleres.


SYSTEMATICS 43<br />

SPICULES. The subtylostyles can be readily divided into two categories by their thickness.<br />

(a) The echinators and some <strong>of</strong> the scattered megascleres are stout, with dimensions<br />

<strong>of</strong> 72-702 X 5-<strong>20</strong> /A. A slight but definite constriction separates the rounded head from<br />

the shaft. The longer spicules are usually curved toward the basal end. The shorter<br />

spicules are usually straight and frequently have roughened heads. A very few stout<br />

subtylostyles exhibit a slight tendency toward shaft spination. The spines, when present,<br />

I L<br />

Fig. 8. —Spicules <strong>of</strong> Microciona rarispinosa. A. Stout subtylostyles. B. Thin subtylostyle. C.<br />

Heads <strong>of</strong> stout tylostyles enlarged to show spination. D. Palmate isochelas. E. Toxa.<br />

A, B: scale I. C, D: scale II. E: scale III. YPM 5041. Holotype.<br />

are only 1-3 in number and usually less than 1 p in height, (b) Thin, scattered subtylostyles,<br />

with narrow ovoid heads, 116-608 X 2-5 p. (c) Narrow palmate isochelas,<br />

12-<strong>20</strong> p long, with a maximum head width <strong>of</strong> 3 fi. (d) Toxas 16-23 X 2 fi or less. Most<br />

are tricurvate, with a wide deep arch, and recurved ends. The smaller toxas have a shallow<br />

arch and straight ends. Individual analyses, lengths in microns:<br />

Thick<br />

Tylostyles<br />

128-702 (50)<br />

72-398 (50)<br />

82-538 (75)*<br />

Thin<br />

Tylostyles<br />

153-375 (50)<br />

116-348 (50)<br />

146-608 (50)*<br />

Isochelas<br />

13-16 (50)<br />

12-15 (25)<br />

13-<strong>20</strong> (25)*<br />

Toxas<br />

16-60 (50)<br />

<strong>20</strong>-44 (25)<br />

<strong>20</strong>-63 (50)*<br />

DISCUSSION. The species is characterized by a lack <strong>of</strong> acanthostyles, and the near absence<br />

<strong>of</strong> shaft spination on the echinating megascleres. It is therefore intermediate between<br />

Microciona and the ophlitaspongiid genus Axociella. In spicule size ranges the present<br />

species is very similar to the Puerto Rican sponges described as Microciona spinosa by<br />

Wilson (1902a). Wilson's species differs by its complete lack <strong>of</strong> spination and by the<br />

presence <strong>of</strong> conules. De Laubenfels (1936a, p. 113) transferred M. spinosa to Axociella.<br />

De Laubenfels' Florida specimens differ from Wilson's in shape, spicule complement and<br />

megasclere distribution. They are therefore unlikely to be conspecific with either the<br />

Puerto Rican or Jamaican sponges.<br />

Microciona similis Stephens (1915) <strong>of</strong> South Africa is also similar in structure to the<br />

Jamaican species but differs in having thicker fibers and larger toxas. The smallest styles<br />

have a considerable number <strong>of</strong> spines on the shaft. De Laubenfels (1936a) established a


44 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

new genus, Axocielita, for M. similis on the erroneous assumption that all its megascleres<br />

are smooth. Axocielita is therefore a synonym <strong>of</strong> Microciona.<br />

The species conforming to de Laubenfels' definition <strong>of</strong> Axocielita can be included<br />

within Axociella Hallmann. De Laubenfels used Axociella for massive sponges with the<br />

same spicule complement as the incrusting species placed in Axocielita. The species<br />

which must be transferred or returned to Axociella include Axociella calla de Laubenfels,<br />

1934; Axocielita kilauea de Laubenfels, 1951a; Axocielita linda de Laubenfels, 1954a;<br />

and Ophlitaspongia membranacea Thiele, 1905. Microciona aceratoobtusa Carter, 1887,<br />

has styles with microspined heads, and can best be retained in Microciona. The presence<br />

<strong>of</strong> a graded series <strong>of</strong> intermediates between Microciona and Axociella suggests that the<br />

validity <strong>of</strong> de Laubenfels* family Ophlitaspongiidae is open to question.<br />

GENUS THALYSEURYPON de Laubenfels<br />

Thalyseurypon conulosa n. sp.<br />

(Text-fig. 9; plate VI, fig. 1)<br />

Fig. 9. —Spicules <strong>of</strong> Thalyseurypon conulosa. A. Subtylostyles. B. Acanthostyle. YPM 5042.<br />

Holotype.<br />

HOLOTYPE. YPM 5042. Off Maiden Cay, August 12, 1961.<br />

HABITAT. The single specimen was collected in about 10 feet <strong>of</strong> water near Maiden<br />

Cay.<br />

SHAPE. The lobate sponge arises from a narrow region <strong>of</strong> attachment. It attains a<br />

maximum height <strong>of</strong> 8 cm and a maximum overall width <strong>of</strong> 10 cm. The partially fused<br />

lobes are 1-2 cm in diameter and several centimeters in height. The sponge is a dark<br />

purple with a reddish tinge in life and retains that color in alcohol. The sponge emits a<br />

purplish exudate and darkens the preservative.<br />

CONSISTENCY. Compressible, but tough and difficult to tear.<br />

SURFACE. The surface is strongly conulose. The conules are 3-6 mm in height, frequently<br />

0.5 mm in thickness, and <strong>of</strong>ten bifid. Adjacent conules are 2-5 mm apart and<br />

are <strong>of</strong>ten connected by thick ridges. A thin, smooth, detachable skin covers both the<br />

conules and interconular areas. The sparsely scattered oscules are less than 0.5 mm in<br />

diameter.<br />

ECTOSOME. The skin is almost entirely aspiculous. Thin oxeote spicules were observed<br />

very rarely in strips <strong>of</strong> the skin. The skin contains extremely abundant oval pigmented<br />

cells, 10-13 ix in diameter. A few possible dermal pores, 21-35 ^ in diameter, were<br />

observed.<br />

ENDOSOME. The skeleton is a network <strong>of</strong> stout spongin fibers with coring and echinating<br />

spicules. The fibers are 40-150 ^ in diameter, mostly 75-175 p apart. The fibers<br />

are composed largely <strong>of</strong> spongin, with only 1-3 rows <strong>of</strong> coring spicules. In places the<br />

skeleton consists <strong>of</strong> single spicules within a thin sheath <strong>of</strong> spongin. All <strong>of</strong> the definitely<br />

coring spicules are smooth. The echinating spicules are <strong>of</strong>ten basally covered with


SYSTEMATICS 45<br />

spongin. Many echinators are entirely smooth. Others have a smooth shaft and finely<br />

spined head. Here and there, an entirely spined acanthostyle can be observed. A very<br />

few thin oxeote spicules are present in the flesh.<br />

SPICULES, (a) Styles, slightly curved to straight, <strong>20</strong>3-449 x 7-10 ^ (125 spicules).<br />

The spicules are somewhat subtylostylote, with a slight subterminal expansion and a<br />

rounded end. For example, a spicule with a shaft diameter <strong>of</strong> 10 microns has a maximum<br />

head diameter <strong>of</strong> 12 p. Below the head, the spicules taper gradually to a sharp point. A<br />

few spicules are definitely styles. Tylostrongyles occur in small numbers. Some echinators<br />

have roughened or finely spined heads. Immature spicules range in form from thin lobate<br />

tylostyles to thin blunt-ended styles. A few spicules <strong>of</strong> typical shape are extremely long.<br />

Spicules <strong>of</strong> 807, 898, 1037 and 1334 ^ were recorded, (b) Echinating acanthostyles, 130-<br />

230 X 5-12 [JL (25 spicules). The very fine spines are scattered irregularly over the entire<br />

surface <strong>of</strong> the spicule. They are 1 /x or less in height, and 2-15 ^ apart, (c) Thin, fusiform<br />

oxeas, 341-652 X 2-3 p. (15 spicules). They are uncommon, but apparently proper<br />

to the sponge.<br />

DISCUSSION. The large size, lack <strong>of</strong> microscleres, and presence <strong>of</strong> some spiny echinators,<br />

places the specimen in the genus Thalyseurypon. The presence <strong>of</strong> some entirely smooth<br />

echinators is noteworthy for the genus. The sponge is quite different from any <strong>of</strong> the<br />

West Indian members <strong>of</strong> the genus. T. vasiformis de Laubenfels (1953a) is a vase with<br />

very large tylostyles. T. foliacea (Topsent, 1889) has spinous coring spicules and leaf-like<br />

branches. T. carteri Topsent (1889) has thick, flat branches, and numerous interstitial<br />

spicules.<br />

Thalyseurypon conulosa is similar in shape, conule size and color to Pandaros acanthifolium<br />

Duchassaing and Michelotti (1864, p. 90). The latter species was described from<br />

St. Thomas and reported from the Dry Tortugas by de Laubenfels (1936a, p. 123).<br />

De Laubenfels reported "occasional acanthose modifications" in the spicules <strong>of</strong> the<br />

St. Thomas specimen (p. 124). T. conulosa differs from Pandaros acanthifolium in having<br />

definite echinators, including a distinctive category <strong>of</strong> small acanthostyles. De Laubenfels<br />

emphasized the fact that "The fibers are not at all echinated" in his redefinition <strong>of</strong><br />

Pandaros (p. 123). T. conulosa also lacks the subdermal spaces and unusually spongy<br />

consistency <strong>of</strong> de Laubenfels* material.<br />

FAMILY MYCALIDAE Lundbeck, 1905 (=Amphilectidae, de Laubenfels 1936a)<br />

A reticulate skeleton. The fibers or spicule tracts are never echinated. The megascleres<br />

are monactinal. The microscleres, usually present, may include sigmas, chelas,<br />

toxas or raphides.<br />

De Laubenfels (1936a) curiously assigned Mycale and Zygomycale to his family<br />

Ophlitaspongiidae, which is defined as having smooth, echinating spicules. The two<br />

genera lack echinators and thus key out to his family Amphilectidae.<br />

Mycale (along with its synonym Esperella, and the preoccupied Esperia) has always<br />

been used for sponges with compact spicule tracts. Topsent (1924), revising the European<br />

species <strong>of</strong> Mycale, described the skeleton as consisting <strong>of</strong> polyspicular fibers. The writer<br />

has found no mention <strong>of</strong> echinators in any description <strong>of</strong> mycalid species, including<br />

those <strong>of</strong> de Laubenfels. Mycale has indeed been used by many authors as the type genus<br />

<strong>of</strong> a family characterized by the lack <strong>of</strong> echinators.<br />

Gray (1867, p. 531) established a family Esperiadae to include Esperia and Mycale.<br />

(The two genera were distinguished on the basis <strong>of</strong> their microsclere complement).<br />

Vosmaer (1886, p. 353) noted that Esperia was a preoccupied name and replaced it by<br />

Esperella. Ridley and Dendy (1887), divided their family Desmacidonidae into two subfamilies,<br />

the Esperellinae and the Ectyoninae. The former subfamily was characterized<br />

by non-echinated, and the latter by echinated fibers. Thiele (1905, p. 949) placed Esperella<br />

into synonymy with Mycale, and Lundbeck (1905, p. 7) accordingly replaced the<br />

name Esperellinae with Mycalinae. The group was raised to a family in the classification<br />

<strong>of</strong> Hentschel (1923).<br />

The Ophlitaspongiidae and Amphilectidae, as listed by de Laubenfels (1936a) probably<br />

require further revision, but many <strong>of</strong> the genera are unfamiliar to me.


46 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

GENUS MYCALE Gray<br />

Mycale laevis (Carter) de Laubenfels<br />

(Plate VI, fig. 2)<br />

Esperia laevis Carter, 1882, p. 291 [type: Puerto Cabello, Venezuela; Brit. Mus. (Nat.<br />

Hist.) (?)]<br />

Mycale laevis, de Laubenfels, 1936a, p. 116.<br />

HABITAT. Very common on pilings at Port Royal.<br />

SHAPE. A typical specimen is fist-sized, massive and lobate. Cylindrical specimens also<br />

occur. An example growing on a crab's back is a horizontally spreading mass, two mm<br />

to 2 cm in thickness.<br />

COLOR. In life, the exterior is a light orange, while the interior is a dull yellow. A<br />

specimen dried after initial preservation in alcohol became white in color. In alcohol,<br />

the sponges are beige, with or without traces <strong>of</strong> the original orange color.<br />

CONSISTENCY. Spongy, compressible.<br />

SURFACE. Smooth to the touch, but uneven in places. The oscules are 0.5-1.5 cm in<br />

diameter. Each receives the openings <strong>of</strong> several large exhalant canals. In some cases, a<br />

deep cloaca is present. The oscules are usually apical on the lobes. They are bordered by<br />

a very characteristic membranous rim which is sometimes divided into filiform processes.<br />

The resultant collar may reach a centimeter in height.<br />

ECTOSOME. The oscular rim contains stout, longitudinal, occasionally anastomosing<br />

spicule bundles, 100-600 fx in diameter. They are also joined by connective tracts. The<br />

rim membranes contain numerous microscleres, including rosettes <strong>of</strong> large anisochelas.<br />

A detachable skin (which easily becomes macerated) covers the surface. The dermal<br />

skeleton includes an irregular reticulation <strong>of</strong> spicule bundles, 40-145 jx in diameter. The<br />

bundles grade into loosely organized groups <strong>of</strong> spicules. A considerable number <strong>of</strong><br />

megascleres are strewn in the flesh. The interstices <strong>of</strong> the network are <strong>of</strong>ten 70-150 fx in<br />

width. They contain microscleres <strong>of</strong> all types, except for large anisochelas. The scattered<br />

ostia are 30-145 ju, in diameter. Their partition walls are <strong>of</strong>ten 30-40 ju. in width and contain<br />

relatively few microscleres (mostly raphides and small anisochelas.)<br />

ENDOSOME. The interior contains scattered megascleres and a coarse irregular network<br />

<strong>of</strong> spicule bundles. The interstitial spicules are <strong>of</strong> varying thickness, while those<br />

<strong>of</strong> the tracts are robust. The reticulate skeleton becomes more prominent near the<br />

periphery. Numerous tracts ascend to the surface, sometimes branching in the process,<br />

to end below the dermis in slightly expanded tufts. Conspicuous rosettes <strong>of</strong> a dozen or<br />

more anisochelas are common near the ascending tracts. The periphery <strong>of</strong> the endosome<br />

also contains a great abundance <strong>of</strong> very compact trichodragmata, 50-70 /*, in length and<br />

about 10 fx in diameter. Such compact bundles are <strong>of</strong>ten clustered together. Compact<br />

trichodragmata, in smaller numbers, occur throughout the sponge along with single<br />

raphides and loose sheaves <strong>of</strong> raphides similar to those figured by Carter. Sigmas are<br />

present in great numbers in the flesh.<br />

SPICULES, (a) Styles, sometimes slightly subtylostylote, straight to somewhat curved,<br />

with a narrowed base, and rather abrupt point, 377-596 x 7-18 p. (b) Large palmate<br />

anisochelas, with a straight, thick shaft, 57-100 JX total length x 7-10 [x maximum shaft<br />

diameter. The large (upper) end consists <strong>of</strong> a prominent wide central tooth flanked by<br />

attached palms, similar in length to the tooth. The lower end is expanded transversely,<br />

terminating at each side in a plate which is directed away from the shaft on the same<br />

side <strong>of</strong> the spicule as the central upper tooth. The upper end <strong>of</strong> the spicule is about as<br />

long as the tooth gap and is between two and three times the length <strong>of</strong> the lower end.<br />

The respective figures for a typical chela are 34, 30, and 13 [i. The maximum width <strong>of</strong><br />

the upper end is usually about 25-40 LI. The lower end is 2/3 to % the width <strong>of</strong> the larger<br />

end. (c) Small anisochelas, with very long upper ends, and narrow, short, lower ends,<br />

15-28 jx total length X 2 /x shaft diameter. As seen in pr<strong>of</strong>ile, the upper tooth extends<br />

down nearly to the level <strong>of</strong> the lower end. The lower end is sometimes extended by a<br />

small apical projection about 2 LL in length, (d) Sigmas, 17-77 X 1-3 p. (e) Raphides,


SYSTEMATICS 47<br />

27-102 X 2 /i. Many are grouped into the previously described trichodragmata and<br />

sheaves. Individual analyses, lengths in microns, 50 styles and 25 <strong>of</strong> each category <strong>of</strong><br />

microsclere per sponge:<br />

Styles<br />

463-565<br />

421-541<br />

467-596<br />

492-590<br />

377-536<br />

Large<br />

Anisochelas<br />

68-87<br />

65-86<br />

70-96<br />

57-100<br />

63-82<br />

Small<br />

Anisochelas<br />

15-23<br />

17-28<br />

17-23<br />

<strong>20</strong>-27<br />

17-22<br />

Sigmas<br />

18-50<br />

17-63<br />

18-57<br />

23-77<br />

24-46<br />

Raphides<br />

50-87<br />

48-102<br />

51-85<br />

27-93<br />

50-87<br />

DISTRIBUTION. Tropical Atlantic America: Venezuela—Carter, 1882, p. 291 (as Esperia<br />

laevis).<br />

DISCUSSION. The Jamaican sponge resembles Carter's Venezuelan species in form,<br />

spicule assortment and spicule sizes. The shapes <strong>of</strong> the spicules are, for the most part, in<br />

close agreement. The small anisochelas <strong>of</strong> my specimens, however, have at best a slight<br />

lower projection which never attains the proportions shown in Carter's figures. Carter<br />

mentions neither membranous nor feathery rims for the oscules. They are certainly quite<br />

characteristic <strong>of</strong> the Jamaican sponge. Being delicate, they may have been destroyed on<br />

the specimens available to Carter.<br />

Mycale microsigmatosa (Arndt) Burton<br />

Mycale fistulata var. microsigmatosa Arndt, 1927, p. 144 [type: Curasao; repository not<br />

specified.]<br />

Mycale microsigmatosa, Burton, 1956, p. 129.<br />

Mycale cecilia de Laubenfels, 1936b, p. 447; idem, 1950b, p. 24; Wells et al., 1960, p.<br />

212 (?)<br />

Mycale senegalensis LeVi, 1952, p. 46; idem, 1959, p. 129.<br />

HABITAT. Common on mangrove roots and on turtle grass.<br />

SHAPE. All <strong>of</strong> the dozens <strong>of</strong> specimens seen in the field were incrustations, 1-3 mm in<br />

thickness.<br />

COLOR. In life, pale orange or pink, or most frequently a dull bluish-green. Bright<br />

red-orange specks, presumably embryos, are present in most specimens at all times <strong>of</strong> the<br />

year. The sponges are predominantly beige in alcohol.<br />

CONSISTENCY. S<strong>of</strong>t.<br />

SURFACE. Smooth to the touch, even. In life, the oscules have delicate, transparent,<br />

raised collars. The oscules are supplied by river-like systems <strong>of</strong> subdermal exhalant canals.<br />

The canals are <strong>of</strong>ten several millimeters in diameter and extend outward for as much<br />

as 1 cm.<br />

ECTOSOME. A thin, detachable dermal membrane is pierced by numerous ostia, 35-<br />

1<strong>20</strong> JU, in diameter. The partitions between adjacent pores are 2-5 p, in thickness and<br />

contain scattered microscleres. The pores are clustered into groups <strong>of</strong> 3-12 or more by<br />

dense bands <strong>of</strong> flesh, which are 10-30 //, in diameter. The skin is supported by the plumose<br />

ends <strong>of</strong> ascending tracts, but megascleres are not present in the dermal membrane.<br />

ENDOSOME. The skeletal framework consists <strong>of</strong> spicule tracts <strong>20</strong>-100 p in diameter. The<br />

tracts occasionally branch or anastomose in the interior. They <strong>of</strong>ten subdivide in ascending<br />

to the surface. Microscleres are abundant throughout the sponge. The anisochelas are<br />

not localized by size. The ovoid embryos are 175-350 ^ in longest diameter.<br />

SPICULES, (a) Tylostyles, straight to curved, with narrow, somewhat ovoid heads and<br />

hastate points, 214-304 X 3-12 /A. The maximum diameter is at the mid-length <strong>of</strong> the<br />

shaft, (b) Sigmas, strongly curved, 30-50 x 1-3 /x. (c) Anisochelas, narrow and palmate,<br />

15-27 p. The shaft is about 2 /x in diameter. The upper palm has a maximum width <strong>of</strong><br />

7 ft in larger spicules. It is usually equal to i/2 or more <strong>of</strong> the total length <strong>of</strong> the spicule.


48 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

Some <strong>of</strong> the smaller spicules have very short shafts. Individual analyses, lengths in microns,<br />

50 megascleres each, and 25 microscleres per category each:<br />

Tylostyles<br />

214-287<br />

234-275<br />

234-264<br />

228-287<br />

234-304<br />

Sigmas<br />

33-46<br />

30-42<br />

33-50<br />

30-44<br />

33-46<br />

Anisochelas<br />

16-25<br />

16-23<br />

17-27<br />

15-26<br />

15-23<br />

DISTRIBUTION. Tropical Atlantic America: North Carolina—Wells et al., 1960, p. 212<br />

(as Mycale cecilia) ?; Curasao, Arndt, 1927, p. 144 (as Mycale fistulata var. microsigmatosa).<br />

Tropical West Africa: Senegal—LeVi, 1952, p. 46 (as Mycale senegalensis). Hawaiian<br />

Is.—de Laubenfels, 1950b, p. 24 (as Mycale cecilia). Tropical Pacific America:<br />

Pacific coast <strong>of</strong> Panama—de Laubenfels, 1936b, p. 447 (as Mycale cecilia).<br />

DISCUSSION. The anisochelas were <strong>of</strong> a single size range in all <strong>of</strong> the Jamaican sponges.<br />

Continuous ranges have also been reported by Arndt (1927), de Laubenfels (1950b)<br />

and LeVi (1952). A spicule preparation obtained from one <strong>of</strong> de Laubenfels' Hawaiian<br />

specimens (U. S. National <strong>Museum</strong> number 22747) contains anisochelas 13-24 ^ in length.<br />

Wells et al., (1960) reported two distinct sizes <strong>of</strong> anisochelas. Their North Carolina material<br />

differs further from the Jamaican in having tangential dermal tylostyles and larger<br />

sigmas. The dermal membrane <strong>of</strong> USNM specimen 22747 contains only microscleres.<br />

De Laubenfels (1936b) reported two sizes <strong>of</strong> anisochelae in his original description <strong>of</strong><br />

Mycale cecilia but noted the presence <strong>of</strong> intermediates in the legend for figure 41.<br />

LeVi (1959) has already suggested that Mycale senegalensis L£vi, 1952 might be conspecific<br />

with Arndt's M. fistulata microsigmatosa. The Curacao sponge is certainly not<br />

conspecific with the tubular Australian Mycale fistulata Hentschel, 1911. M. cecilia de<br />

Laubenfels cannot be distinguished from Arndt's variety and must be regarded as a<br />

synonym <strong>of</strong> M. microsigmatosa.<br />

Closely related species are known from Hawaii (M. maunakea de Laubenfels, 1951a)<br />

and Australia (M. phyllophila Hentschel, 1911). M. maunakea differs from M. microsigmatosa<br />

in having unusually narrow isochelas. Its spicule tracts neither branch nor<br />

anastomose. M. phyllophila, while very similar to the West Indian sponge, is geographically<br />

remote. M. microsigmatosa is therefore retained as a separate species.<br />

GENUS ZYGOMYCALE Topsent<br />

Zygomycale parishii (Bowerbank) Topsent<br />

(Plate V, fig. 3)<br />

Raphiodesma parishii Bowerbank, 1875, p. 283 [type: Straits <strong>of</strong> Malacca; Brit. Mus. (Nat.<br />

Hist.) (?)]<br />

Esperia parishii, Ridley, 1884, p. 436.<br />

Esperella parishii, Ridley and Dendy, 1887, p. 65; Dendy, 1905, p. 159.<br />

Zygomycale parishii, Topsent, 1929, p. 431; de Laubenfels, 1950b, p. 25; idem, 1956, p.<br />

3; Levi, 1956a, p. 14.<br />

Mycale parishii, Burton and Rao, 1932, p. 328.<br />

Esperia plumosa Carter, 1882, p. 298 (fide Burton and Rao, 1932); idem, 1887, p. 72.<br />

Esperella plumosa, Dendy, 1905, p. 159; idem, 1916a, p. 121; Dendy and Frederick, 1924,<br />

p. 503.<br />

Esperella ridleyi Lendenfeld, 1888, p. 211 (fide Burton and Rao, 1932); Hallmann, 1914,<br />

p. 402.<br />

Mycale isochela Hentschel, 1911, p. 297 (fide Burton and Rao, 1932.)<br />

Mycale pectinicola Hentschel, 1911, p. 299 (fide Burton and Rao, 1932.)<br />

HABITAT. Specimens were collected on pilings at Port Royal and also at Gunboat<br />

Beach on the peninsula between Port Royal and Kingston.


SYSTEMATICS 49<br />

SHAPE. The sponge has a small, incrusting or irregularly massive base from which<br />

arise long, subdividing, anastomosing branches. Many specimens reach <strong>20</strong>-25 cm in length.<br />

The branches are usually only a few millimeters in diameter but occasionally reach a<br />

width <strong>of</strong> 1.5-2 cm.<br />

COLOR. In life the peripheral layer may be tan, brown, grayish-brown, or purple.<br />

Toward the interior the color gradually becomes a dull yellow. In alcohol, the sponges<br />

may be pale yellow, beige, or partially purple.<br />

CONSISTENCY. Somewhat compressible but tough.<br />

SURFACE. The sponge is covered by a skin which is smooth to the touch. The overall<br />

appearance is uneven. In addition to numerous projections and branches, the skin is<br />

thrown into low elevations, ridges and conules. On the branches, the conules are sometimes<br />

several millimeters in height. The small scattered oscules are less than 1 mm in<br />

diameter.<br />

ECTOSOME. A pore-bearing membrane is supported by a reticulation <strong>of</strong> loose to compact<br />

spicule tracts, which are 30-50 p in diameter. The meshes are triangular, irregularly polygonal,<br />

or roughly rectangular in outline. Parallel bundles are frequently 150-250 p apart.<br />

The interstices are crowded with dermal pores separated by narrow walls <strong>of</strong>ten only 2-3 JX<br />

in width. Typical ostial sizes are 40 x 30, 70 x 60, and 88 X 40 ju. The partition walls<br />

contain infrequent microscleres, with isochelas being the most numerous category. Rosettes<br />

<strong>of</strong> large anisochelas are frequently present near the bundles. Interstitial megascleres are<br />

infrequent.<br />

ENDOSOME. The interior is microcavernous, particularly near the periphery. Subdermal<br />

spaces are occasionally present. The skeleton consists <strong>of</strong> a very irregular network <strong>of</strong> spicule<br />

tracts. The tracts are 40-<strong>20</strong>0 p, in diameter. Spongin is sometimes present. At the periphery,<br />

tracts ascend to the surface, to end in slightly expanded tufts under the dermal membrane.<br />

Many megascleres are strewn in the flesh. Conspicuous rosettes <strong>of</strong> 6-10 large<br />

anisochelas are present in the periphery. The endosome contains an abundance <strong>of</strong><br />

microscleres, with isochelas the most numerous, and large sigmas the least numerous<br />

category. Small sigmas may be very abundant or quite infrequent. Aspiculous ovoid<br />

embryos, about 300 X 400 p as seen in section, were found in specimens collected in<br />

February and June.<br />

SPICULES, (a) Subtylostyles, occasionally becoming styles, with narrow elongate heads,<br />

curved to undulating shafts, and gradually narrowing apices, 217-334 x 3-12 /x. The<br />

maximum diameter is at the mid-length <strong>of</strong> the shaft. A few megascleres have abnormal<br />

swellings and projections, (b) Large palmate anisochelas, 37-67 X 2 p shaft diameter.<br />

They are similar in shape to the anisochelas <strong>of</strong> Mycale laevis. The width <strong>of</strong> the upper<br />

end is 15-<strong>20</strong> p; that <strong>of</strong> the lower end 10-12 p. A typical spicule 48 microns long has an<br />

upper tooth <strong>of</strong> 24 p, a lower end <strong>of</strong> 9 p and a tooth gap <strong>of</strong> 15 p. The spicules occur<br />

singly or, more frequently, in rosettes near the periphery <strong>of</strong> the endosome. (c) Small<br />

palmate anisochelas, 13-28 tt long. The maximum width at the upper end is about 6 /*,.<br />

A spicule 17 p long has an upper tooth <strong>of</strong> 10 p, a lower end <strong>of</strong> 4 p, and tooth gap <strong>of</strong> 3 ju,.<br />

(d) Palmate isochelas, 10-13 p long. The cap-shaped ends are about 3/x in width. Very<br />

abundant, (e) Large sigmas, with sharply recurved ends, 54-106 x 2-7 p.. They are very<br />

abundant, (f) Small slender sigmas, 15-42 X 1-2 p. Varied in abundance, (g) Toxas, <strong>of</strong>ten<br />

in toxodragmata <strong>of</strong> several spicules, 22-107 X 2 p. They have a shallow, gradually curved<br />

arch, and flattened to slightly recurved ends, (h) Raphides, <strong>of</strong>ten in trichodragmata, 23-<br />

53 X 1-2 JU,. All categories are present in the seven analysed specimens. Individual analyses,<br />

lengths in microns, 50 megascleres per sponge, 25 microscleres <strong>of</strong> each category per<br />

sponge:<br />

Large Small<br />

Styles Anisochelas Anisochelas Isochelas<br />

236-326 43-67 13-28 10-13<br />

234-334 46-54 16-28 10-13<br />

217-314 45-52 17-22 10-13<br />

239-290 37-52 17-<strong>20</strong> 10-12


50 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

Large<br />

Sigmas<br />

61-91<br />

88-106<br />

54r-90<br />

54-85<br />

Small<br />

Sigmas<br />

<strong>20</strong>-33<br />

16-42<br />

16-34<br />

15-34<br />

Toxas<br />

28-80<br />

33-107<br />

27-77<br />

22-89<br />

Raphides<br />

23-53<br />

30-40<br />

24-44<br />

24-34<br />

DISTRIBUTION. Tropical Atlantic America: Brazil—de Laubenfels, 1956, p. 3. Indo-<br />

Pacific: Madagascar—L£vi, 1956a, p. 14; Mauritius, Carter, 1882, p. 298 (as Esperia<br />

plumosa); Okhamandal, India—Dendy, 1916a, p. 121 (as Esperella plumosa); Ceylon—<br />

Dendy, 1905, p. 159 (as Esperella parishii); Dendy, 1905, p. 159 (as Esperella plumosa);<br />

Mergui Archipelago, <strong>of</strong>f Burma—Carter, 1887, p. 72 (as Esperia plumosa); Malaya—<br />

Bowerbank, 1875, p. 283 (as Raphiodesma parishii); Burton and Rao, 1932, p. 328 (as<br />

Mycale parishii); Philippines—Ridley and Dendy, 1887, p. 65 (as Esperella parishii);<br />

Australia—Ridley, 1884, p. 436 (as Esperia parishii); Lendenfeld, 1888, p. 211 (as Esperella<br />

ridleyi), Hentschel, 1911, p. 297 (as Mycale isochela), Hentschel, 1911, p. 299 (as Mycale<br />

pectinicola); Dendy and Frederick, 1924, p. 503 (as Esperella plumosa); Hawaiian Is.—<br />

de Laubenfels, 1950b, p. 25.<br />

DISCUSSION. The Jamaican specimens are the first West Indian record <strong>of</strong> Zygomycale<br />

parishii. Except for de Laubenfels* specimen from Brazil, all previous records have come<br />

from the Pacific and Indian Oceans.<br />

De Laubenfels (1950b) and Burton and Rao (1932), have concluded that all <strong>of</strong> the<br />

species included in the taxonomic summary are conspecific with the type-species Zygomycale<br />

parishii (Bowerbank). Their conclusions are probably correct, as all <strong>of</strong> the supposed<br />

species are certainly very similar in appearance and architecture. Zygomycale parishii has<br />

seven types <strong>of</strong> microscleres. The other species are, in large part, distinguished only by the<br />

lack <strong>of</strong> one or more <strong>of</strong> those spicule types. In size and form, their microscleres are similar<br />

to the corresponding ones <strong>of</strong> Z. parishii. Burton and Rao (1932) concluded from the<br />

literature and their own material that Z. parishii is a highly variable species, in which<br />

individuals <strong>of</strong>ten lacked one or more <strong>of</strong> the seven kinds <strong>of</strong> microsclere. De Laubenfels,<br />

however, reported no such variation in his Hawaiian specimens, nor did mine vary in<br />

spicule complement.<br />

A review <strong>of</strong> the literature suggests that the apparent differences between specimens<br />

and supposed species may be due to microsclere categories being overlooked or taken for<br />

developmental stages. Ridley (1884), for example, considered small anisochelas and sigmas<br />

as the early stages <strong>of</strong> the larger spicules. Dendy (1916a) noted that he had apparently<br />

made the same assumption in his earlier discussions <strong>of</strong> Esperella plumosa. Spicule types<br />

have certainly been overlooked in many studies. Hallmann found several additional<br />

categories in the type <strong>of</strong> Esperella ridleyi Lendenfeld. He found no trace <strong>of</strong> the rare<br />

diancistras attributed to the sponge by Lendenfeld.<br />

Esperia plumosa was established by Carter (1882) for a specimen from Mauritius<br />

which had isochelas, two (?) sizes <strong>of</strong> anisochela, one size <strong>of</strong> sigma, and toxas. In 1887,<br />

Carter reported a Burmese specimen which had trichodragmata in addition to the spicules<br />

<strong>of</strong> the Mauritius sponge. Dendy (1905) examined Carter's type and additional specimens<br />

from Ceylon. He recorded toxodragmata (but not trichodragmata or small anisochelas)<br />

in all the specimens. In 1916, as mentioned above, he amended his earlier account by<br />

listing small sigmas and anisochelas for his specimens from India and also for the<br />

earlier specimens from Ceylon, and Carter's type. He concluded (1916a, p. 122) that<br />

"the constancy in form and size in all the specimens I have examined, including the<br />

type, is very remarkable." Burton and Rao, however, were unable to find rosettes in one<br />

<strong>of</strong> Dendy's specimens and toxas in another. They did find previously unrecorded trichodragmata<br />

in his specimens. Adding positive reports, we find that Dendy's examples<br />

have all seven kinds <strong>of</strong> microsclere, and Carter's type at least six.<br />

Bowerbank's Z. parishii, although the type species, is poorly known. His small specimen<br />

was apparently growing over a myxillid, and the spicules <strong>of</strong> both were included<br />

in the original description (cf. Ridley 1884). Ridley concluded, presumably incorrectly,


SYSTEMATICS 51<br />

that the toxas mentioned by Bowerbank belonged to the myxillid. He and Dendy (1887)<br />

attributed a Philippine specimen to parishii on the assumption that neither had toxas.<br />

Upon re-examination, Dendy (1905) found toxas in the Challenger specimen.<br />

Mycale pectinicola and M. isochela Hentschel differ from a typical Z. parishii only<br />

by the apparent absence <strong>of</strong> toxas in the first species, and <strong>of</strong> the larger forms <strong>of</strong> chelas<br />

and sigmas in the second. Hentschel (1911, p. 301) indeed suspected that all the isochelate<br />

mycalids known to him were conspecific.<br />

Mycale crassissima (Dendy, 1905) is considered as a synonymy <strong>of</strong> Z. parishii by de<br />

Laubenfels (1950b, p. 27). The original description, which erroneously included isochelas,<br />

has been corrected by Hentschel (1912) and Dendy (1916b, p. 55).<br />

GENUS ULOSA de Laubenfels<br />

Ulosa hispida n. sp.<br />

(Text-fig. 10)<br />

Fig. 10.—Spicules <strong>of</strong> Ulosa hispida. Styles. YPM 5043. Holotype.<br />

HOLOTYPE. YPM 5043. Mangrove boat channels, Feb. 21, 1961.<br />

HABITAT. Common on mangrove roots near Port Royal.<br />

SHAPE. The sponge spreads over mangrove roots as an incrustation which is 1-3 mm<br />

in total thickness. The sponge has a very uneven outline, with numerous low, irregular<br />

projections.<br />

COLOR. The living sponges are a light orange. They fade to a pale beige in alcohol.<br />

CONSISTENCY. S<strong>of</strong>t, slimy, delicate.<br />

SURFACE. The surface is uneven and hispid. Numerous spicules project singly beyond<br />

the surface for most <strong>of</strong> their length. In life, thin delicate collared oscules can be seen.<br />

ECTOSOME. No dermal specialization.<br />

ENDOSOME. The skeleton is a reticulation <strong>of</strong> slender spongin fibers cored by smooth<br />

styles. The fibers are 15-90 /x in diameter and contain only 1-4 spicules at any one place.<br />

The fibers branch acutely and occasionally anastomose. They are frequently joined by<br />

short aspiculous connective bonds, 15-30 p in diameter. The projecting spicules arise<br />

from the fibers. They do not differ in form or size range from those <strong>of</strong> the interior.<br />

Some <strong>of</strong> the projecting spicules are partially or even entirely covered by a thin film<br />

<strong>of</strong> spongin. The flesh contains some spicule debris and a considerable quantity <strong>of</strong> fine<br />

sand. No styles are present in the flesh.<br />

SPICULES. Styles, straight to slightly curved, gradually pointed, 493-1193 X 5-12 /*.<br />

Individual analyses, lengths in microns: 503-1193 (50); 515-1070 (75); 493-898 (50); 562-<br />

1053 (50); 573-971 (50); 507-1135 (50).* A great variety <strong>of</strong> spicules, including oxeas <strong>of</strong> a<br />

wide range <strong>of</strong> sizes, were observed. Most <strong>of</strong> the spicules could easily be referred to common<br />

mangrove species. The spicules were most abundant in spicule preparations for<br />

which mangrove twigs were scraped. Careful study leads to the conclusion that only the<br />

styles are proper to the sponge.<br />

DISCUSSION. The species is provisionally placed in the genus Ulosa, which has been<br />

utilized by de Laubenfels for sponges in which the spicule complement consists only <strong>of</strong><br />

smooth styles. (Cf. Ulosa rhoda de Laubenfels, 1957, and Ulosa spongia de Laubenfels,<br />

1954a). De Laubenfels (1936a, p. 126) originally defined the genus as having coring styles<br />

and a mixture <strong>of</strong> styles and strongyles in the flesh. However, Topsent (1930, p. 27)<br />

redescribed the type-species (Spongia angulosa Lamarck) as having numerous styles, and<br />

a feeble number <strong>of</strong> oxeas. Strongylacidon intermedia Burton (1934, p. 550), with coring<br />

strongyles which are occasionally modified to styles, was also placed in the genus by


52 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

de Laubenfels. Burton's species is probably not congeneric with any <strong>of</strong> the species now in<br />

Ulosa.<br />

The present species differs sharply from the type-species in appearance and spicule<br />

size. It does seem congeneric with U. spongia and U. rhoda de Laubenfels. The distinctive<br />

characters <strong>of</strong> the Jamaican sponge include the large size <strong>of</strong> its spicules, the abundance<br />

<strong>of</strong> foreign matter, the spongin-rich fibers, and the hispidation <strong>of</strong> the surface by single<br />

spicules.<br />

ORDER HALICHONDRIDA Vosmaer 1887, sensu de Laubenfels, 1936a<br />

FAMILY HALICHONDRIIDAE Gray, sensu de Laubenfels, 1936a<br />

GENUS HALICHONDRIA Gray<br />

Halichondria melanadocia de Laubenfels<br />

(Plate V, fig. 2)<br />

Halichondria melanadocia de Laubenfels, 1936a, p. 133 [type: Dry Tortugas; USNM no.<br />

22463]; Wells et al., 1960, p. 225; Little, 1963, p. 54.<br />

HABITAT. Abundant on pilings and mangrove roots near Port Royal.<br />

SHAPE. Specimens on mangrove roots <strong>of</strong>ten spread horizontally along the roots, attaining<br />

a thickness <strong>of</strong> one to several centimeters. Others are irregularly massive, lobate<br />

or even partly ramose.<br />

COLOR. The exterior is grayish-black. The interior has a very characteristic yellowishgreen<br />

tinge. In alcohol, the specimens are dark gray to nearly black with a drab interior.<br />

CONSISTENCY. Compressible.<br />

SURFACE. Smooth to the touch. Projecting lobes and low oscular chimneys <strong>of</strong>ten make<br />

the surface rather uneven. The oscules, which also may be flush with the surface, are<br />

2-3 mm in diameter. They have a membranous, irregularly outlined rim.<br />

ECTOSOME. The detachable dermal skin contains a tangential reticulation <strong>of</strong> compact<br />

spicule bundles, 35-130 p in diameter. The interstices are irregularly polygonal in shape,<br />

and 100-<strong>20</strong>0 ^ in maximum width. They contain a few scattered spicules and one to<br />

several dermal pores. Typical ostial sizes are 40 x 30, 90 X 60, and 145 x 90 p. The<br />

subdermal channels are very extensive and <strong>of</strong>ten reach 1 mm in depth. They open widely<br />

into macroscopic inhalant canals which run toward the interior. The thick partitions<br />

between the subdermal spaces are fleshy extensions <strong>of</strong> the endosome, 300-700 fi in diameter.<br />

They contain scattered spicules and also spicule tracts which run obliquely towards<br />

the surface.<br />

ENDOSOME. The interior is cavernous and confused. The thickly strewn spicules are<br />

occasionally grouped into loosely organized tracts.<br />

SPICULES. Oxeas, straight to considerably curved, 134-802 X 3-<strong>20</strong> p. The ends may be<br />

gradually pointed, somewhat hastate, irregular in outline, or even stair-stepped. The<br />

spicules are not localized in the sponge by size. Individual analyses, lengths in microns,<br />

50 spicules each: 140-656; 164-686; 134-597; 134-691; 141-743; 152-802.<br />

DISTRIBUTION. Tropical Atlantic America: North Carolina—Wells et al., 1960, p. 225<br />

(?); Dry Tortugas, Florida—de Laubenfels, 1936a, p. 133; West Coast, Florida—Little,<br />

1963, p. 54.<br />

DISCUSSION. The dark Jamaican Halichondria agrees closely with H. melanadocia in<br />

structure and external appearance. De Laubenfels' brief description is at least not inconsistent<br />

with the dermal structure found in my specimens. It would be <strong>of</strong> great interest<br />

to know if the tangentially arranged dermal spicules <strong>of</strong> his sponge are merely strewn,<br />

or grouped into bundles. The fetid odor mentioned by de Laubenfels did not occur in<br />

my sponges. A spicule preparation obtained from the type contains oxeas 116-507 fi in<br />

length. Most <strong>of</strong> the spicules are straight and hastate. Their ends are occasionally irregular<br />

or stair-stepped. The Jamaican Halichondria is therefore provisionally considered as<br />

conspecific with that <strong>of</strong> de Laubenfels.<br />

The North Carolina record listed by Wells et al. concerns a thin brown conulose<br />

incrustation. Further specimens will be needed to confirm the identification.


SYSTEMATIGS 58<br />

Halichondria magniconulosa n. sp.<br />

(Text-fig. 11; plate V, fig. 1)<br />

Fig. 11.—Spicules <strong>of</strong> Halichondria magniconulosa. Oxeas. YPM 5039. Holotype.<br />

HOLOTYPE. YPM 5039. Rasta's Wreck, June 24, 1961.<br />

HABITAT. Several specimens were found on pilings at Port Royal.<br />

SHAPE. Massive to lobate, reaching a height <strong>of</strong> as much as 11 cm.<br />

COLOR. The surface and interior <strong>of</strong> the living sponge is yellow. In alcohol, exposed<br />

surfaces become mottled grayish-brown to dark brown. The interior becomes a dull<br />

yellow.<br />

CONSISTENCY. The sponge is tough and only slightly compressible.<br />

SURFACE. The skin is wrinkled, but smooth to the touch. It can be detached only<br />

with difficulty. The surface is frequently thrown into thick, sharp-pointed conules. The<br />

conules are 1-7 mm in height, several millimeters in width at the base, and typically<br />

about 1 mm in thickness. They may be sparsely scattered or spaced only a few millimeters<br />

apart. In the two smaller specimens, the infrequent oscules are 1-3 mm in diameter. The<br />

largest specimen has two apical vents, each being 1 cm in diameter.<br />

ECTOSOME. The dermal membrane receives the terminal tufts <strong>of</strong> radial spicule tracts.<br />

It also contains rather sparsely scattered tangential spicules <strong>of</strong> all sizes. Small spicules are<br />

particularly abundant. The dermal pores range in size from 40 x 30-100 X 60 /x. They<br />

are divided into ill-defined groups by bands <strong>of</strong> flesh which are 15-60 p in diameter. The<br />

spicules within the partitions may be scattered or organized into loose bundles. Some<br />

areas <strong>of</strong> the skin seem to lack pores.<br />

ENDOSOME. The interior skeleton includes a feltwork <strong>of</strong> scattered spicules and also<br />

spicule bundles, 30-100 p in diameter. The well-developed radial tracts end in dermal<br />

tufts. Numerous spicule bundles enter each conule. Subdermal spaces are infrequent.<br />

SPICULES. Oxeas, mostly straight, occasionally slightly curved, 108-369 x 3-7 p. They<br />

may be fusiform or somewhat hastate. Small spicules (123-166 p) and large spicules (290-<br />

340 fi) are particularly common but intermediates do occur. The spicules are not localized<br />

by size within the sponge. Individual analyses, lengths in microns, 50 spicules each: 116-<br />

363*; 108-348; 123-369.<br />

DISCUSSION. The massive shape and thick conules <strong>of</strong> the Jamaican yellow Halichondria<br />

separate it from both H. panicea and H. bowerbanki. The dermis <strong>of</strong> the<br />

Jamaican sponge resembles that <strong>of</strong> H. bowerbanki, but is less well developed. Many <strong>of</strong><br />

the spicules seen in strips <strong>of</strong> the skin are clearly the ends <strong>of</strong> radial bundles. Subdermal<br />

cavities are quite restricted in my specimens. The Jamaican sponges have a much lower<br />

limit <strong>of</strong> spicule size than either North Carolina or New England summer specimens <strong>of</strong><br />

H. bowerbanki. In addition, the smaller spicules are very abundant in my material.<br />

The marked darkening in alcohol is a striking characteristic <strong>of</strong> the new species. H.<br />

bowerbanki, by contrast, becomes a pale yellowish-green in alcohol.


54 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

ORDER HADROMERIDA Topsent, 1898b<br />

FAMILY SPIRASTRELLIDAE Ridley and Dendy, sensu de Laubenfels, 1936a<br />

GENUS SPIRASTRELLA Schmidt<br />

Spirastrella coccinea (Duchassaing and Michelotti) de Laubenfels<br />

Thalysias coccinea Duchassaing and Michelotti, 1864, p. 84 [type: St. Thomas; Brit. Mus.<br />

(Nat. Hist.) reg. no. 28.11.12.46.]<br />

Spirastrella coccinea, de Laubenfels, 1936a, p. 143; idem, 1949, p. 19; idem, 1950a, p. 96;<br />

Dickinson, 1945, p. 36; Wells et al., 1960, p. 228; Little, 1963, p. 55.<br />

Spirastrella cunctatrix Schmidt, 1868, p. 17; Carter, 1879a, p. 357; idem, 1881a, p. 384;<br />

idem, 1882, p. 351; idem, 1886, p. 112 (fide Dendy, 1897); idem, 1887, p. 188; Topsent,<br />

1894a, p. 44; idem, 1918, p. 544; idem, 1925a, p. 630; idem, 1938, p. 21; Dendy, 1897,<br />

p. <strong>20</strong>2, 253; Arndt, 1927, p. 140; Burton, 1936a, p. 22; de Laubenfels, 1936b, p. 461;<br />

idem, 1956, p. 2; LeVi, 1957, p. <strong>20</strong>4; idem, 1959, p. 124; Sara, 1958a, p. 221; idem,<br />

1960a, p. 451; Sara and Siribelli, 1960, p. 39; Sara, 1961a, p. 34.<br />

Chondrilla phyllodes Schmidt, 1870, p. 26 (fide Topsent, 1918); Topsent, 1889, p. 32;<br />

Carter, 1890, p. 565.<br />

Chondrillina phyllodes, de Laubenfels, 1936a, p. 182.<br />

Spirastrella decumbens, Topsent (not Ridley, 1884, p. 470), 1897, p. 440 (fide Topsent,<br />

1918.)<br />

Spirastrella mollis Verrill, 1907, p. 344 (fide de Laubenfels, 1950a.)<br />

Spirastrella purpurea, Vosmaer, 1911a, p. 9, 11 [partim]; idem, 1933, p. 416 [partim];<br />

(not Alcyonium purpureum Lamarck, 1815.)<br />

HABITAT. Drunkenman's Cay; on rocks.<br />

SHAPE. Incrusting, about 1 mm thick.<br />

COLOR. One specimen was dull orange in life and is a pale grayish-brown in alcohol.<br />

The second was brown in life and is a pale brown in preservative.<br />

CONSISTENCY. Tough, slightly compressible.<br />

SURFACE. Smooth to the touch; even to very slightly wrinkled. In the living sponge,<br />

river-like systems <strong>of</strong> subdermal canals lead to small oscules. Neither vents nor canals are<br />

evident in preserved material.<br />

ECTOSOME. No detachable skin. Spirasters, mostly <strong>of</strong> small size, form a cortical layer<br />

40-100 ii in thickness. Bands <strong>of</strong> microscleres inclose dermal pores which are about 30 /*, in<br />

diameter.<br />

ENDOSOME. Numerous tylostyles are strewn in confusion. A few spicule tracts are<br />

present, particularly in the originally brown specimen. The tracts are <strong>of</strong>ten obliquely<br />

oriented. They end in plumose tufts at the surface. Spirasters, mostly <strong>of</strong> large size, are<br />

abundant in the flesh.<br />

SPICULES, (a) Tylostyles, pin-like, with long, straight, gradually pointed shafts and<br />

distinct heads, 281-480 x 3-12 fi (typically 5-9 ^t). The heads may be ovoid, lobate, or<br />

occasionally spherical. They are about 9 p in diameter, (b) Spirasters, prominently spined,<br />

7-58 x 2-7 //,. The microscleres have a wide variety <strong>of</strong> forms, which are all connected<br />

by intermediates. The smaller spicules typically have a slender, slightly curved shaft<br />

which bears spines 2-3 /A in height. Some are amphiasters, with spines grouped at the<br />

ends <strong>of</strong> a straight shaft. Most <strong>of</strong> the larger spicules have a stout curved axis, with one<br />

or two bends. Their conical spines are 5-7 fx or even as much as 13 /x in height. Some<br />

have a rather short axis with strong spines and resemble the spicules <strong>of</strong> Diplastrella<br />

bistellata (Schmidt) Topsent. A few <strong>of</strong> the endosomal spirasters have a long, thin axis<br />

and low spines. They are <strong>20</strong>-25 x 2 fi, and have spines 2 ^ in height. Individual analyses,<br />

lengths in microns, 50 spicules per category:<br />

Tylostyles Spirasters<br />

334-445 7-47<br />

281-480 8-58<br />

DISTRIBUTION. Tropical Atlantic America: North Carolina—Wells et al., 1960, p. 228;<br />

Dry Tortugas, Florida—de Laubenfels, 1936a, p. 143; West Coast, Florida—Little, 1963,


SYSTEMATICS 55<br />

p. 55; Gulf <strong>of</strong> Campeche, Mexico—Topsent, 1889, p. 32 (as Chondrilla phyllodes); Atlantic<br />

coast <strong>of</strong> Panama—de Laubenfels, 1936b, p. 461 (as Spirastrella cunctatrix); Brazil—<br />

de Laubenfels, 1956, p. 2 (as Spirastrella cunctatrix); Bermuda—Verrill, 1907, p. 344 (as<br />

Spirastrella mollis), de Laubenfels, 1950a, p. 96; Bahamas—de Laubenfels, 1949, p. 19;<br />

Antilles—Schmidt, 1870, p. 26 (as Chondrilla phyllodes); St. Thomas, W. I.—Duchassaing<br />

and Michelotti, 1864, p. 84 (as Thalysias coccinea); Curasao—Arndt, 1927, p. 140 (as<br />

Spirastrella cunctatrix); Fernando Noronha, tropical South Atlantic—Carter, 1890, p.<br />

565 (as Chondrilla phyllodes)}<br />

Mediterranean—Atlantic—Ligurean Sea, Italy—Sara, 1958a, p. 221 (as Spirastrella<br />

cunctatrix); Ischia, Italy—Sara, 1960a, p. 451 (as Spirastrella cunctatrix); Naples, Italy—<br />

Sara and Siribelli, 1960, p. 39 (as Spirastrella cunctatrix), Topsent, 1925a, p. 630, (as<br />

Spirastrella cunctatrix); Adriatic Sea—Sara, 1961a, p. 34 (as Spirastrella cunctatrix);<br />

Israel—LeVi, 1957, p. <strong>20</strong>4 (as Spirastrella cunctatrix); Mediterranean <strong>of</strong>f Egypt—Burton,<br />

1936a, p. 22 (as Spirastrella cunctatrix), Tunisia—Topsent, 1894a, p. 44 (as Spirastrella<br />

cunctatrix); Algeria—Schmidt, 1868, p. 17, and Carter, 1882, p. 351 (both as Spirastrella<br />

cunctatrix); Cyprus—Schmidt, 1868, p. 17 (as Spirastrella cunctatrix); West Africa—<br />

Topsent, 1918, p. 544, and LeVi, 1959, p. 124 (both as Spirastrella cunctatrix).<br />

Indo-Pacific: Indian Ocean—Mauritius—Carter, 1879a, p. 357, and 1882, p. 351 (both<br />

as Spirastrella cunctatrix); Mergui Archipelago, <strong>of</strong>f Burma—Carter, 1887, p. 188 (as<br />

Spirastrella cunctatrix).<br />

Pacific Ocean: East Indies—Topsent, 1897, p. 440 (as Spirastrella decumbens) and<br />

Vosmaer, 1911a, p. 9, 11 (as Spirastrella purpurea) partim; Australia—Carter, 1879a, p.<br />

357, 1881a, p. 384, 1882, p. 351 (all as Spirastrella cunctatrix)}.<br />

Tropical Pacific America: Gulf <strong>of</strong> California—Dickinson, 1945, p. 36.<br />

DISCUSSION. The spirasters <strong>of</strong> the Jamaican sponges are similar in range <strong>of</strong> size and<br />

shape to the microscleres figured by Topsent (1918, p. 543, 545) for Spirastrella cunctatrix<br />

Schmidt. Topsent later verified his identification by comparison with Schmidt's type.<br />

(cf. Topsent, 1938). S. cunctatrix, along with its synonym Chondrilla phyllodes Schmidt,<br />

has been recorded from the West Indies by Topsent (1889, 1918) and Arndt (1927).<br />

Thalysias coccinea Duchassaing and Michelotti, 1864, was originally described as a<br />

red incrusting sponge with short-rayed stelliform spicules. De Laubenfels (1936a), who<br />

had apparently examined the type, transferred the species to Spirastrella. He distinguished<br />

S. coccinea from S. cunctatrix largely by the smaller size <strong>of</strong> the spirasters in the<br />

former species, (cf. de Laubenfels, 1956, p. 2). De Laubenfels did, however, report rather<br />

large spirasters in a Bermudan sponge attributed to S. coccinea. (cf. de Laubenfels, 1950a).<br />

An examination <strong>of</strong> material identified as 5. coccinea by de Laubenfels suggests that<br />

S. cunctatrix and S. coccinea are conspecific. The specimens include one from Bermuda<br />

in the <strong>Yale</strong> <strong>Peabody</strong> <strong>Museum</strong> (YPM 714). It contains spirasters 13-34 ^ in length. U. S.<br />

National <strong>Museum</strong> specimen 22498, collected by de Laubenfels in the Dry Tortugas, has<br />

spirasters 10-34 p in length. The two sponges are indistinguishable from Sv cunctatrix<br />

and the Jamaican sponges in architecture and range <strong>of</strong> spiraster form. Most <strong>of</strong> the<br />

spirasters in one Jamaican specimen (having an overall size range <strong>of</strong> 7-47 fx) are 35 ^ or<br />

less in length. The observed differences in maximum length are almost certainly individual<br />

variations. S. cunctatrix is therefore considered to be a synonym <strong>of</strong> the earliernamed<br />

S. coccinea.<br />

Carter's Indo-Pacific records require verification. His material included massive specimens.<br />

Their spirasters, as figured by Vosmaer (1911a), are not dissimilar to those <strong>of</strong><br />

S. cunctatrix. Dendy (1887) considered S. cunctatrix variety robusta Carter, 1886 to be<br />

specifically distinct from the Mediterranean S. cunctatrix. He transferred Carter's variety<br />

porcata (1886) to another species.<br />

GENUS ANTHOSIGMELLA Topsent<br />

Anthosigmella varians (Duchassaing and Michelotti) de Laubenfels<br />

(Plate VII, figs. 4, 5)<br />

Thalysias varians Duchassaing and Michelotti, 1864, p. 86 [type: St. Thomas; Brit. Mus.<br />

(Nat. Hist.) reg. no. 28.11.12.48.]


56 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

Anthosigmella varians, de Laubenfels, 1936a, p. 143; idem, 1949, p. 19; idem, 1953a, p.<br />

539; idem, 1954a, p. <strong>20</strong>3; Little, 1963, p. 55.<br />

[non] Anthosigmella varians, Wells et al., 1960, p. 228.<br />

Suberites tuberculosus Schmidt, 1870, p. 46 (fide Topsent, 19<strong>20</strong>.)<br />

Suberites coronarius Carter (not Carter, 1887, p. 74.), 1882, p. 352 (fide de Laubenfels,<br />

1936a.)<br />

Spirastrella coronaria, Topsent, 1894, p. 26.<br />

Anthosigmella coronarius, Topsent, 1918, p. 557; idem, 19<strong>20</strong>a, p. 30.<br />

[non] Cliona coronaria, Dendy, 1916a, p. 132.<br />

Papillina arcuata Topsent, 1889, p. 35 (fide Topsent, 1894.)<br />

HABITAT. Several specimens were dredged from mussel beds in about 10 feet <strong>of</strong><br />

water near the Kingston side <strong>of</strong> the Port Royal mangrove thickets. One specimen was<br />

collected at a similar depth near the cays.<br />

SHAPE. Irregularly massive.<br />

COLOR. The upper surface is brownish, the lower one cream to brown. The interior<br />

is a drab yellow. In alcohol, the sponges are beige with yellow, green or rust-red tinges.<br />

CONSISTENCY. Somewhat rubbery, but rather stiff.<br />

SURFACE. The sponge may have occasional irregular ridges and projections. Most <strong>of</strong><br />

the surface is covered by small, flattened, welt-like tubercles, about 1 mm in diameter.<br />

At a magnification <strong>of</strong> 60 x > the surface is hispid with projecting spicules. Small oscules<br />

were detected in one specimen.<br />

ECTOSOME. The deeply pigmented periphery is packed with tylostyles, mostly with<br />

points directed outward.<br />

ENDOSOME. Microcavernous. The megascleres are thickly strewn around the canals.<br />

Occasional subdermal spaces, 175-500 p, deep, are present under the dermal palisade.<br />

SPICULES, (a) Tylostyles with well-developed oblong heads, slightly constricted necks<br />

and somewhat curved shafts, tapering to a point, 194-467 X 3-18 /x. (b) Anthosigmas,<br />

with knob-like spines on the ends and convex side <strong>of</strong> a slightly to strongly curved shaft,<br />

13-25 x 2 p. A few spicules are amphiasters, and a very few tend toward the spirastral<br />

form. The microscleres are abundant in all specimens. Individual analyses, lengths in<br />

microns:<br />

Tylostyles Anthosigmas<br />

269-451 (70) 13-25 (50)<br />

217-433 (50) 15-25 (50)<br />

19^-467 (50) 15-25 (50)<br />

182-439 (50) 15-23 (50)<br />

DISTRIBUTION. Tropical Atlantic America: Florida—Schmidt, 1870, p. 46 (as Suberites<br />

tuberculosus); de Laubenfels, 1949, p. 19; West Coast, Florida—de Laubenfels, 1953a,<br />

p. 539; West Coast, Florida—Little, 1963, p. 55; Campeche, Mexico—Topsent, 1889, p.<br />

35 (as Papillina arcuata); Honduras—Carter, 1882, p. 352 (as Suberites coronarius)) Bahamas—Carter,<br />

1882, p. 352 (as Suberites coronarius)) de Laubenfels, 1949, p. 19; Jamaica—Carter,<br />

1882, p. 352, (as Suberites coronarius); Lesser Antilles—Guadeloupe, St.<br />

Barthelemy, St. Thomas, Tortole—Duchassaing and Michelotti, 1864, p. 86 (as Thalysias<br />

varians).<br />

DISCUSSION. De Laubenfels (1953a, p. 539-540) has reported considerable variation in<br />

the presence or absence <strong>of</strong> microscleres and in the relative abundance <strong>of</strong> anthosigmas<br />

and typical spirasters. None <strong>of</strong> the other investigators <strong>of</strong> the species have reported such<br />

variation. If all <strong>of</strong> de Laubenfels' specimens are conspecific, the species presents us with<br />

an interesting intermediate between the spirastrellids and the suberitids. Wells et al.<br />

(1960) have assigned very thin incrustations devoid <strong>of</strong> microscleres to A. varians. It is far<br />

more likely that they are a species <strong>of</strong> Prosuberites.<br />

Anthosigmella varians has had a complicated taxonomic history. It was first recognizably<br />

described by Carter. Topsent transferred it to Spirastrella in 1894 on the basis


SYSTEMATIGS 57<br />

<strong>of</strong> the presence <strong>of</strong> microscleres. He concluded that his Papillina arcuata <strong>of</strong> 1889 was<br />

conspecific with Carter's species. In 1918 he established the genus Anthosigmella for<br />

spirastrellids with the peculiar microscleres designated as anthosigmas. De Laubenfels<br />

(1936a, p. 143) concluded, presumably after examining Duchassaing and Michelotti's<br />

type, that Carter's sponge was conspecific with the earlier Thalysias varians. The sponge<br />

figured on plate 17 <strong>of</strong> Duchassaing and Michelotti's paper could well be the present<br />

species. Finally, Topsent (19<strong>20</strong>a) discovered anthosigmas in the type <strong>of</strong> Schmidt's<br />

Suberites tuberculosus and placed it into synonymy with Carter's species.<br />

Thiele (1902) and Annandale (1915) have shown that Carter's Burmese sponge (1887)<br />

is a Cliona. Dendy's Indian record is probably the same species, Cliona orientalis<br />

Thiele, 1903.<br />

GENUS SPHECIOSPONGIA Marshall<br />

Spheciospongia vesparia (Lamarck) Marshall<br />

Alcyonium vesparium Lamarck, 1815, p. 78 [type: Antilles; Mus. nat. Hist. nat. Paris.]<br />

Thalysias vesparia, Duchassaing and Michelotti, 1864, p. 85.<br />

Spheciospongia vesparia, Marshall, 1892, p. 32; de Laubenfels, 1932b, p. 50; idem, 1936a,<br />

p. 140; idem, 1947, p. 34; idem, 1949, p. 18; idem, 1953a, p. 536; Topsent, 1933a, p.<br />

30; Wells et al., 1960, p. 227; Little, 1963, p. 55.<br />

Papillina cribrosa Schmidt, 1870, p. 48 (fide de Laubenfels, 1932b.)<br />

Hymeniacidon pulvinatus Bowerbank, 1872, p. 126 (fide de Laubenfels, 1932b.)<br />

Spirastrella pulvinata, Ridley, 1884a, p. 187.<br />

[non] Spirastrella pulvinata, Arndt, 1927, p. 139.<br />

Spongia dysoni Carter, 1879a, p. 348 (fide de Laubenfels, 1932b); idem, 1882, p. 350.<br />

Spirastrella purpurea, Vosmaer, 1911a, p. 6 [partim.]<br />

Spirastrella andrewsii George and Wilson, 1919, p. 135 (fide de Laubenfels, 1932b.)<br />

Poterion atlantica George and Wilson, 1919, p. 139 (fide de Laubenfels, 1932b.)<br />

HABITAT. The single specimen was collected by Dr. T. F. Goreau <strong>of</strong> the <strong>University</strong><br />

<strong>of</strong> the West Indies in <strong>20</strong> feet <strong>of</strong> water <strong>of</strong>fshore from Port Royal. He has seen numerous<br />

specimens in deeper Jamaican waters.<br />

SHAPE. A large, massive cake-shaped sponge, about one foot in diameter. Only a few<br />

small pieces <strong>of</strong> the sponge were successfully preserved.<br />

COLOR. The living sponge is entirely black. In alcohol, the fragments become gray.<br />

CONSISTENCY. Almost woody when first collected. The fragments are rather tough and<br />

incompressible.<br />

SURFACE. Uneven, with many low, gradually rising elevations. The large oscules (8<br />

and 15 mm wide in the fragments) are not raised above the surface. Numerous openings,<br />

1-1.5 mm in diameter, are scattered singly, and also arranged in sieve areas. Each sieve<br />

has about <strong>20</strong>-25 openings. No microscopic pores could be detected.<br />

ECTOSOME. The cortex is a thick feltwork <strong>of</strong> tylostyles and overlies extensive subdermal<br />

spaces. Spirasters are abundant.<br />

ENDOSOME. Very cavernous, with the smooth-lined canals <strong>of</strong>ten reaching 1 mm in<br />

diameter. The partition walls contain a thick feltwork <strong>of</strong> tylostyles. Microscleres are common<br />

only in the canal linings.<br />

SPICULES, (a) Tylostyles, with small, nearly circular heads, straight to considerably<br />

curved shafts, and gradually narrowing points, 156-472 X 5-17 ^. (75 spicules), (b)<br />

Spirasters, occasionally without bends, but usually zigzagged with 1-6 (typically 3-4) bends<br />

in the shaft, 12-30 ^ in length x 2-3 //, in total diameter. (50 spicules). The numerous low<br />

spines, no more than 2 /x in height, are arranged spirally in at least the larger spicules.<br />

The spines are most prominent at the apices <strong>of</strong> the bends.<br />

DISTRIBUTION. Tropical Atlantic America: North Carolina—George and Wilson, 1919,<br />

p. 135 (as Spirastrella andrewsi) and p. 139 (as Poterion atlantica), de Laubenfels, 1947,<br />

p. 34, Wells et al., 1960, p. 227. Florida—Schmidt, 1870, p. 48 (as Papillina cribrosa),<br />

Dry Tortugas, Florida—de Laubenfels, 1936a, p. 140; West Coast, Florida—de Laubenfels,<br />

1953a, p. 536 and Little, 1963, p. 55; British Honduras—Bowerbank, 1872, p. 126 (as


58 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

Hymeniacidon pulvinatus), Carter, 1879a, p. 348 (as Spongia dysoni); Bahamas—de Laubenfels,<br />

1949, p. 18; Jamaica—George and Wilson, 1919, p. 135 (as Spirastrella andrewsi);<br />

Guadeloupe, St. Croix—Duchassaing and Michelotti, 1864, p. 85 (as Thalysias vesparia).<br />

DISCUSSION. The specimens <strong>of</strong> Lamarck and Schmidt have been reexamined by both<br />

Topsent (1933a) and de Laubenfels (1936a). The two species <strong>of</strong> George and Wilson have<br />

been examined by de Laubenfels and by Wells et al. These authors presumably have<br />

found microscleres in Poterion atlantica. Carter's Spongia dysoni Bowerbank apparently<br />

refers to the sponge which Bowerbank called Hymeniacidon pulvinatus. Carter<br />

took the name from a label in the British <strong>Museum</strong>, but Bowerbank never used it in his<br />

publications. The Curacao sponge assigned to Bowerbank's species by Arndt (1927) is<br />

a Spirastrella with two sizes <strong>of</strong> spirasters.<br />

De Laubenfels considered the Bermudan Spheciospongia as a distinct species, S. othella.<br />

It has enormous raised oscules, patches <strong>of</strong> yellow both on the surface and in the interior,<br />

and somewhat smaller tylostyles than S. vesparia. De Laubenfels has, however, reported<br />

oscules <strong>of</strong> as much as 7 cm in diameter (raised?) in Bahaman examples <strong>of</strong> S. vesparia.<br />

In 1953, he reported an entirely yellow Spheciospongia with a spiculation "exactly that<br />

<strong>of</strong> 5. vesparia." (1953a, p. 537). More specimens will be needed to clarify the number <strong>of</strong><br />

species <strong>of</strong> Spheciospongia in the West Indies.<br />

GENUS DIPLASTRELLA Topsent<br />

Diplastrella megastellata n. sp.<br />

Fig. 12.—Spicules <strong>of</strong> Diplastrella megastellata. A. Tylostyles. B. Two forms <strong>of</strong> spherasters. C.<br />

Small asters. B, C: scale I. A: scale II. YPM 5044. Holotype.<br />

HOLOTYPE. YPM 5044. Maiden Cay, Aug. 12, 1961.<br />

HABITAT. The single specimen was collected at Maiden Cay in a few feet <strong>of</strong> water.<br />

SHAPE. A horizontally spreading crust with a maximum thickness <strong>of</strong> 1 mm. The sponge<br />

covered approximately a square foot <strong>of</strong> the coral substratum.


SYSTEMATICS 59<br />

COLOR. The sponge is a dull brownish olive and retains that color in preservative.<br />

CONSISTENCY. Slightly compressible.<br />

SURFACE. Even. Under a magnification <strong>of</strong> 14 x, the surface appears hispid and slightly<br />

uneven. No oscules could be detected.<br />

ECTOSOME. Little specialization. The spirasters are densely crowded in a peripheral<br />

layer <strong>of</strong> the sponge, 70-150 ^ in thickness. Many tylostyles project beyond the flesh.<br />

ENDOSOME. In many places, only a single layer <strong>of</strong> tylostyles is present in the sponge.<br />

Most <strong>of</strong> the megascleres are erect. Spirasters and their derivatives are numerous throughout<br />

the sponge. They gradually become more abundant toward the surface. A thin layer,<br />

<strong>of</strong>ten a single row, <strong>of</strong> spherasters covers the substratum. The spherastral layer is present<br />

in all parts <strong>of</strong> the sponge. The flesh contains conspicuous spherical cells 7 LI in diameter.<br />

SPICULES, (a) Tylostyles, robust, straight to very slightly curved, 253-841 x 9-26 /x<br />

(range <strong>of</strong> length <strong>of</strong> 100 spicules). The prominent rounded heads are wider than long,<br />

and their diameter exceeds that <strong>of</strong> the shaft, (b) Asters, <strong>of</strong> varied form, 9-<strong>20</strong> LL in total<br />

diameter (50 spicules). The most common form has 4-5 branching rays arising from an<br />

unexpanded central region. Many spicules are spirasters with short, slightly curved,<br />

entirely spined shafts. Terminally spined amphiasters also occur. A few large spicules<br />

are nearly spherastral, and resemble the microscleres described for Diplastrella by<br />

Topsent (1918). The spines and rays are 1-7 /x in height, (c) Spherasters, 29-79 fx in<br />

diameter (50 spicules). Smaller spicules have short simple rays <strong>of</strong> equal length. Larger<br />

spherasters have rays <strong>of</strong> unequal length and varied form. Their shorter rays are usually<br />

blunt-tipped. Their longer rays have an irregular outline, and may be extended into<br />

several blunt, roughened knobs or projections. The rays sometimes resemble an arm with<br />

fingers. The spicules are limited to a basal crust.<br />

DISCUSSION. The present species is closely related to Diplastrella ornata Rutzler and<br />

Sara (1962), described from the Mediterranean coast <strong>of</strong> Italy, and the Adriatic. Both<br />

species have large branching spherasters concentrated in a layer next to the substratum.<br />

The Jamaican sponge lacks the small oxeas found in the European species.<br />

FAMILY SUBERITIDAE Schmidt, sensu de Laubenfels, 1936a<br />

GENUS TERPIOS Duchassaing and Michelotti<br />

Terpios zeteki (de Laubenfels) de Laubenfels<br />

(Plate VII, fig. 3)<br />

Laxosuberites zeteki de Laubenfels, 1936b, p. 450 [type: Balboa, Canal Zone; USNM no.<br />

22212]; Dickinson, 1945, p. 37.<br />

Terpios zeteki, de Laubenfels, 1950a, p. 106; idem, 1950b, p. 28; idem, 1951a, p. 265;<br />

idem, 1954b, p. 339.<br />

HABITAT. The species is abundant on mangrove roots along the larger channels.<br />

SHAPE. Large, massive, with lobate or digitate projections. The branches are <strong>of</strong>ten<br />

10 cm in height and 1 cm in diameter. Specimens are <strong>of</strong>ten considerably larger than<br />

hand-sized. Dickinson has aptly described the sponge as "a mass <strong>of</strong> giant fingers fused."<br />

(1945, p. 37).<br />

COLOR. The interior and the basal surfaces are yellowish-orange. The remainder <strong>of</strong><br />

the surface is one <strong>of</strong> a wide range <strong>of</strong> colors, including red, red-orange, orange, green,<br />

blue-green and lavender. The color is always uniform within a specimen. Reddish-brown<br />

and blue specimens have been recorded in the literature. The sponges are light gray in<br />

alcohol.<br />

CONSISTENCY. Tough, slightly compressible.<br />

SURFACE. Basically smooth to the touch, and even. The sponge is, however, quite contractile.<br />

Preserved specimens, as well as many in the field, may thus appear wrinkled and<br />

tuberculate. At a magnification <strong>of</strong> 60 X, the surface is microhispid. The oscules are <strong>of</strong>ten<br />

as large as a centimeter in the living animal but are usually quite obscure, due to contraction,<br />

in preserved specimens.<br />

ECTOSOME. The external color is confined to a thin peripheral layer. Large dermal<br />

tufts form a nearly continuous palisade at the surface. The spicules project beyond the


60 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

surface for y3 to i/2 <strong>of</strong> their length, with their apices nearly on a level over considerable<br />

distances. The radiating tufts have broad bases. Tylostyles <strong>of</strong> all sizes are found in the<br />

tufts.<br />

ENDOSOME. The tylostyles, again <strong>of</strong> all sizes, are strewn abundantly in the flesh. They<br />

are grouped into bundles near the periphery. While the overall pattern is one <strong>of</strong> confusion,<br />

the spicules tend to be oriented longitudinally in the axis <strong>of</strong> the surface projections.<br />

SPICULES. Tylostyles, straight, fusiform, gradually pointed, 140-877 X 3-18 /x. The<br />

prominent heads are typically lobate but occasionally rounded. Individual analyses,<br />

lengths in microns, 50 spicules each. The color <strong>of</strong> each specimen is indicated. 164-819<br />

(red-orange); 134-726 (orange); 152-796 (green); 170-749 (red); 140-877 (orange); 164-<br />

644 (lavender); 143-819 (orange); 140-842 (orange); 170-830 (orange).<br />

DISTRIBUTION. Tropical Atlantic America: Atlantic coast <strong>of</strong> Panama—de Laubenfels,<br />

1936b, p. 450 (as Laxosuberites zeteki). Indo-Pacific: Hawaiian Is.—de Laubenfels, 1950b,<br />

p. 28, 1951a, p. 265, 1954b, p. 339. Tropical Pacific America: Gulf <strong>of</strong> California—Dickinson,<br />

1945, p. 37 (as Laxosuberites zeteki); Pacific coast <strong>of</strong> Panama—de Laubenfels,<br />

1936b, p. 450 (as Laxosuberites zeteki).<br />

DISCUSSION. The present record extends the distribution <strong>of</strong> Terpios zeteki into the<br />

Antilles. The color variants are identical in structure and spiculation. A variety <strong>of</strong> colors,<br />

a wide range <strong>of</strong> spicule length, and a large maximum spicule size, are all characteristics<br />

<strong>of</strong> the species.<br />

FAMILY CLIONIDAE Gray, sensu de Laubenfels, 1936a<br />

GENUS CLIONA Gray<br />

Cliona vermifera Hancock<br />

Cliona vermifera Hancock, 1867, p. 239 [type: locality unknown; Brit. Mus. (Nat. Hist.)<br />

(?)]; Topsent, 1889, p. 35; idem, 1900, p. 8; idem, 1933b, p. 565; Vosmaer, 1933, p.<br />

398; Volz, 1939, p. 18.<br />

Vioa vermifera, Lendenfeld, 1897, p. 80.<br />

HABITAT. The specimen was found in coral rubble at Maiden Cay in a few feet <strong>of</strong><br />

water.<br />

SHAPE. Scattered papillae, 0.5-1.5 mm in width, extend to the surface <strong>of</strong> the coral.<br />

The interior canals are 1-3 mm in diameter.<br />

COLOR. In life, bright red. The preserved sponge is a dull yellow.<br />

STRUCTURE. The canal linings contain numerous tylostyles strewn in confusion. A considerable<br />

number <strong>of</strong> microscleres are present. Granular, spherical cells, 7-12 fi in diameter,<br />

are abundant and conspicuous.<br />

SPICULES, (a) Tylostyles, straight to curved with prominent heads and gradually<br />

pointed shafts, 137-391 x 5-10 /x (100 spicules). The heads are 7-13 fi in maximum diameter.<br />

Most <strong>of</strong> them are round to lobate, but some are subterminal. Polytylote spicules are<br />

rare. The shaft exhibits little or no constriction below the head. The longer megascleres<br />

are mostly straight and rather thin. Their shaft diameter is no greater than, and may<br />

be less than, that <strong>of</strong> the smaller spicules, (b) Spirasters, smooth, curved to undulating,<br />

with rounded ends, 32-70 X 2-5 fi. (50 spicules). The diameter is uniform along the<br />

length <strong>of</strong> most spicules. A few spirasters have a swollen or lumpy region, which, when<br />

present, is near the midlength <strong>of</strong> the spicule. The two ends <strong>of</strong> a spicule may lie along<br />

a single axis or be considerably out <strong>of</strong> line. The curves <strong>of</strong> a spicule may vary in amplitude<br />

along its length. Most spicules have from two to five bends, but some are nearly<br />

straight, and others have as many as seven turns.<br />

DISTRIBUTION. Tropical Atlantic America: Gulf <strong>of</strong> Campeche, Mexico—Topsent, 1889,<br />

p. 35. Mediterranean-Atlantic: Naples, Italy—Vosmaer, 1933, p. 565; Adriatic Sea—<br />

Lendenfeld, 1897, p. 80, (as Vioa vermifera), Volz, 1939, p. 18.<br />

Indo-Pacific: Tuamotu and Gambier Archipelagos, Central Pacific—Topsent, 1933b,<br />

p. 565.<br />

DISCUSSION. The species has been only infrequently reported, although its geographic


SYSTEMATICS 61<br />

range is extensive. The spirasters <strong>of</strong> the Adriatic specimens are seldom, if ever, multiply<br />

bent.<br />

Cliona viridis (Schmidt) Gray<br />

Vioa viridis Schmidt, 1862, p. 77 [type: Zara Channel, Yugoslavia; repository unknown];<br />

Lendenfeld, 1897, p. 59; Burton, 1936a, p. 21.<br />

Cliona viridis, Gray, 1867, p. 545; Topsent, 1896, p. 123; idem, 1900, p. 84; idem, 1902b,<br />

p. 345; idem, 1906a, p. 564; idem, 1925a, p. 630; idem, 1925b, p. 18; idem, 1928, p.<br />

146; idem, 1933b, p. 562; idem, 1934, p. 13; Annandale, 1915, p. 13; Kumar, 1925,<br />

p. 228; Burton, 1930, p. 496; idem, 1948, p. 755; Volz, 1939, p. 13; LeVi, 1957, p. <strong>20</strong>2;<br />

Sara and Siribelli, 1960, p. 39; Sara, 1961a, p. 38; Sara, 1961b, p. 14; Vacelet, 1960, p.<br />

261; idem, 1961, p. 30.<br />

Cliona viridis var. caribboea, Topsent, 1933b, p. 563.<br />

[non] Cliona viridis, Wells et al., 1960, p. 32; Little, 1963, p. 58.<br />

Papillina nigricans Schmidt, 1862, p. 69 (fide Topsent, 1900); idem, 1868, p. 15.<br />

Osculina polystomella Schmidt, 1868, p. 3 (fide Topsent, 1900).<br />

Hymeniacidon angulata Bowerbank, 1872, p. 632 (fide Topsent, 1902b).<br />

Cliona subulata Sollas, 1878, p. 65 (fide Topsent, 1900); Topsent, 1889, p. 35.<br />

Cribrella labiata Keller, 1880, p. 275 (fide Topsent, 1900).<br />

Cliona caribboea Carter, 1882, p. 346 (fide Topsent, 1900); Topsent, 1889, p. 49.<br />

Cliona celata, Vosmaer (not Grant, 1826), 1933, p. 349 [partim.]<br />

HABITAT. TWO specimens were collected at Maiden Cay, boring in coral rubble.<br />

SHAPE. The low surface papillae are 1-1.5 mm in diameter. The galleries are 1-3 mm<br />

in diameter.<br />

COLOR. The papillae are yellow in life and dull yellow-green in alcohol.<br />

STRUCTURE. The canal linings contain thickly strewn tylostyles and numerous spirasters.<br />

Oval refractile cells, 7-10 p diameter, are also abundant. The papillae contain a single<br />

rank <strong>of</strong> parallel tylostyles. The infrequent spirasters <strong>of</strong> the papillae are similar in size<br />

and shape to those <strong>of</strong> the galleries.<br />

SPICULES, (a) Tylostyles, mostly straight, sometimes slightly curved, 195-384 x 5-12 ju,.<br />

The maximum shaft diameter is exceeded by that <strong>of</strong> the head. The spherical to ovoid<br />

heads are usually 13-15 p, in diameter. The maximum head diameter is almost always<br />

near the terminus. A low second swelling may be present just below the head. The shaft<br />

gradually diminishes to a sharp point, (b) Spirasters, thin, covered with spines, typically<br />

multiangular, <strong>20</strong>-48 p in length, with a shaft diameter <strong>of</strong> 1-2 p. The numerous spines<br />

are thin, conical and 2-7 p in length. They are most prominent on the convex parts <strong>of</strong><br />

the shaft. Most spicules have 3-4 bends, but the range is 1-7. A very few spicules are<br />

straight or nearly so. Individual analyses, lengths in microns:<br />

Tylostyles Spirasters<br />

239-384 (125) 24-48 (75)<br />

195-398 (50) <strong>20</strong>-42 (50)<br />

DISTRIBUTION. Tropical Atlantic America: Gulf <strong>of</strong> Campeche, Mexico—Topsent, 1889,<br />

p. 35 (as Cliona subulata); Lesser Antilles: Guadeloupe—Topsent, 1889, p. 49 (as Cliona<br />

caribboea); St. Vincent—Carter, 1882, p. 346 (as Cliona caribboea); European Boreal<br />

Atlantic: Norway—Burton, 1930, p. 496. Mediterranean-Atlantic: French Mediterranean<br />

—Topsent, 1896, p. 123; 1900, p. 84; 1925b, p. 18; Monaco—Topsent, 1934, p. 13; Naples,<br />

Italy—Topsent, 1925a, p. 630; Sara and Siribelli, 1960, p. 39; Sara, 1961b, p. 14; Capri,<br />

Italy—Keller, 1880, p. 275 (as Cribrella labiata); Adriatic—Schmidt, 1862, p. 77 (as Vioa<br />

viridis); Schmidt, 1862, p. 69 (as Papillina nigricans); Lendenfeld, 1897, p. 59 (as Vioa<br />

viridis); Volz, 1939, p. 13; Sara, 1961a, p. 38; Israel—LeVi, 1957, p. <strong>20</strong>2; Mediterranean<br />

coast <strong>of</strong> Egypt—Burton, 1936a, p. 21 (as Vioa viridis); Algeria—Schmidt, 1868, p. 15 (as<br />

Papillina nigricans); 1868, p. 3 (as Osculina polystomella); Topsent, 1902b, p. 345;<br />

Corsica—Vacelet, 1960, p. 261 and 1961, p. 30; Madeira Is.—Bowerbank, 1872, p. 632, (as


62 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

Hymeniacidon angulata); Topsent, 1928, p. 146; Cape Verde Is.—Topsent, 1928, p. 146;<br />

Congo coast—Burton, 1948, p. 755.<br />

Indo-Pacific: Red Sea—Topsent, 1906a, p. 564; India—Kumar, 1925, p. 228; Margui<br />

Archipelago, <strong>of</strong>f Burma—Annandale, 1915, p. 13; Polynesia—Topsent, 1933b, p. 562<br />

(including Cliona viridis caribboea).<br />

DISCUSSION. Clionids with robust, large-headed tylostyles and long, multiangular,<br />

prominently spined spirasters have been recorded from many warm water areas. Spiraster<br />

thickness varies in specimens from the same locality, (cf. Topsent 1900, 1933b). Several<br />

species may be involved, but the Cliona viridis complex cannot be subdivided on present<br />

evidence. European specimens do tend to have a larger maximum tylostyle size than<br />

West Indian specimens. The Jamaican sponges have smaller papillae than most but not<br />

all European specimens, (cf. Topsent 1900, Volz 1939). Mediterranean sponges exhibit<br />

a strong tendency toward the fusion <strong>of</strong> papillae and for development into the massive<br />

form. (Topsent, 1900; Volz 1939; Hartman, personal communication). Studies on growth<br />

may provide a basis for distinguishing the West Indian sponges from C. viridis <strong>of</strong> Mediterranean<br />

waters.<br />

Sponges <strong>of</strong> the C. viridis type have been recorded from the West Indies as C. caribboea<br />

Carter (by Carter, 1882, and Topsent, 1889) and as C. subulata Sollas (by Topsent,<br />

1889). The spicule sizes <strong>of</strong> the Jamaican sponges are in close agreement with the descriptions<br />

<strong>of</strong> Carter and Topsent. (cf. Topsent, 1900, for spicule data on his 1889 material).<br />

Wells et al. (1960) listed Suberites undulatus George and Wilson, 1919, as a synonym <strong>of</strong><br />

C. viridis. Dr. W. D. Hartman has informed me that this species is not a clionid and that<br />

George and Wilson's name is valid. Sponges without microscleres have been attributed<br />

to C. caribboea Carter by Verrill (1907), de Laubenfels (1936a, 1936b, 1950a, 1953a), and<br />

Wells et al. (1960). The relationship <strong>of</strong> their sponges to the spiraster-containing West<br />

Indian clionids is uncertain.<br />

FAMILY PLACOSPONGIIDAE Gray, 1867<br />

GENUS PLACOSPONGIA Gray<br />

Placospongia carinata (Bowerbank) Carter<br />

(Plate VII, fig. 1)<br />

Geodia carinata Bowerbank, 1858a, p. 308, 314 (manuscript name); idem, 1864, p. 254,<br />

pi. X (manuscript name); idem, 1874a, p. 298 [type: South Seas; Brit. Mus. (Nat<br />

Hist.) (?)]; idem, 1875b, p. 295.<br />

Placospongia carinata, Carter, 1879a, p. 148; Ridley, 1884, p. 481; Sollas, 1888, p. 272;<br />

Lindgren, 1897, p. 485; idem, 1898, p. 45; Vosmaer and Vernhout, 1902a, p. 3; Dendy,<br />

1905, p. 126; idem, 1916a, p. 132; idem, 1921, p. 144; Hentschel, 1912, p. 324; Levi,<br />

1956a, p. 10; Burton, 1959a, p. 213; Little, 1963, p. 56.<br />

Placospongia sp., Carter, 1880, p. 53.<br />

Placospongia intermedia Sollas, 1888, p. 272 (fide Vosmaer and Vernhout, 1902a); de<br />

Laubenfels, 1936b, p. 454.<br />

Placospongia mixta Thiele, 1900, p. 72 (fide Vosmaer and Vernhout, 1902a.)<br />

HABITAT. The sponge grows over hard objects near Port Royal and at the cays.<br />

SHAPE. Incrusting. Branched specimens are known for the species, but none were seen<br />

in the present study. Specimens several square feet in area were observed on the sea<br />

wall <strong>of</strong> the Police Depot at Port Royal. None <strong>of</strong> them were more than a few millimeters<br />

in thickness.<br />

COLOR. The cortical plates are brown in life, with red or orange tinges. The flesh<br />

is yellow and retains a drab yellow color in preservative. The surface becomes a very<br />

uniform chocolate brown in alcohol. Dried specimens are pale gray.<br />

CONSISTENCY. The flesh is rather tough. The cortical plates are stony.<br />

SURFACE. The surface is covered by an armor <strong>of</strong> cortical plates, separated by a network<br />

<strong>of</strong> grooves. The yellow flesh extends to the surface at the grooves, which are open as


SYSTEMATICS 63<br />

much as 1 mm in life. Delicate cylindrical oscules were observed in the grooves in uncontracted<br />

living specimens. The plates became apposed in specimens removed from the<br />

water. The plates are irregularly polygonal in shape and have ridged edges. Their outlines<br />

may be straight, curved or irregular. Parallel sides are 2-12 mm apart.<br />

ECTOSOME. An ectochrote <strong>of</strong> microstrongyles covers the plates and grooves. The plates,<br />

<strong>of</strong>ten 1 mm in thickness, are dense aggregations <strong>of</strong> mature sterrasters. The ascending<br />

columns <strong>of</strong> flesh contain microstrongyles and spir asters. Laterally placed, projecting<br />

bundles <strong>of</strong> tylostyles extend into the plates and grooves from the endosome. Throughout<br />

their length the bundles are inclosed by fibrous tissue which also binds adjacent plates<br />

together. The grooves are crossed by vertical canals which are surrounded by microscleres<br />

and presumably open at the surface.<br />

ENDOSOME. The interior contains stout bundles <strong>of</strong> tylostyles which begin near a basal<br />

layer <strong>of</strong> sterrasters and penetrate the cortex. The bundles gradually expand to a maximum<br />

in the mid-endosome where they are <strong>of</strong>ten 300 ^ in diameter. The tylostyles are<br />

oriented obliquely inward in the bundles, with their apices directed toward the periphery.<br />

A few tylostyles occur outside the bundles. They are mostly tangential in position at the<br />

periphery <strong>of</strong> the endosome. The flesh contains numerous sterrasters at all stages <strong>of</strong><br />

development and scattered spirasters. Microstrongyles are common only in the outer<br />

part <strong>of</strong> the endosome and above the basal layer <strong>of</strong> sterrasters. In places, the flesh may<br />

be packed with sterrasters.<br />

SPICULES, (a) Tylostyles, straight with fusiform shafts, 164-1193 X 3-17 /x. Smaller<br />

spicules have sharp points, but larger ones are <strong>of</strong>ten tylostrongylote with narrow, rounded<br />

points. The heads are typically round and prominent but are occasionally elongate or<br />

even subterminal.<br />

(b) Sterrasters, ovoid but indented at the hilum, 36 x 14 to 87 X 73 /*,. The mature<br />

spicules are <strong>of</strong>ten about 75 X 60 /x. A developmental sequence similar to that presented<br />

in Figure 9 <strong>of</strong> Vosmaer and Vernhout (1902a) can be followed in the endosomal sterrasters.<br />

The earliest recognizable stage is about 30 x 7-8 ju, and consists <strong>of</strong> a straight,<br />

narrow spinous shaft. The next stage is a cylindrical spicule, about 35 x <strong>20</strong> ti, with<br />

numerous spines which are 3-5 //, in height. It is succeeded by a spiny, rather dumbbellshaped<br />

spicule <strong>of</strong> 40 X 25 p. The final developmental stage has a round outline with<br />

low spines.<br />

(c) Spirasters, 13-30 x 2-3 fx. The spicules may be spirastral, streptastral or amphiastral<br />

in form. The spines, 3-9 ju, in height, may be scattered along a curved or multiangular<br />

shaft, or be concentrated terminally on a straight shaft. They are <strong>of</strong>ten branched<br />

or terminally bifid. They are usually slender, pointed and irregularly curved or bent.<br />

A few spicules have short shafts and resemble the arborescent-elongo-subsphero-stellate<br />

spicule figured by Bowerbank (1858a, 1864). All variations can occur within a specimen.<br />

(d) Microstrongyles, straight to curved or multiangular, roughened, with rounded<br />

ends, 7-17 X 2 /x.<br />

(e) A few spherasters were found in the spicule analyses <strong>of</strong> two specimens. The spicules<br />

are 21-29 fi in diameter and have short, conical spines. As Vosmaer and Vernhout<br />

have shown, the spherasters are infrequent and, indeed, are <strong>of</strong>ten absent, in both Placospongia<br />

carinata and P. melobesioides.<br />

Individual analyses, lengths in microns:<br />

Tylostyles<br />

240-1193 (50)<br />

164-1005 (50)<br />

229-819 (50)<br />

Microstrongyles<br />

10-15 (25)<br />

7-12 (25)<br />

8-17 (25)<br />

Sterrasters<br />

41 X 23-82 X 64 (50)<br />

47 X 27-87 X 73 (50)<br />

36xl4r-79x57(50)<br />

Spherasters<br />

28(1)<br />

none<br />

21-29 (30)<br />

Spirasters<br />

13-30 (25)<br />

13-30 (25)<br />

15-30 (25)


64 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

DISTRIBUTION. Tropical Atlantic America: West Coast, Florida—Little, 1963, p. 56;<br />

Atlantic coast <strong>of</strong> Panama—de Laubenfels, 1936b, p. 454 (as Placospongia intermedia))<br />

Indo-Pacific: Madagascar—LeVi, 1956a, p. 10; South Arabian coast—Burton, 1959a, p.<br />

213; Indian Ocean, south <strong>of</strong> Seychelles—Dendy, 1921, p. 144; Okhamandal, northwest<br />

India—Dendy, 1916a, p. 132; Ceylon—Dendy, 1905, p. 125; Straits <strong>of</strong> Malacca—Bowerbank,<br />

1875b, p. 295 (as Geodia carinata); South Seas, Bowerbank—1858a, p. 308, 314;<br />

1864, p. 254; 1874a, p. 289 (all as Geodia carinata); East Indies—Vosmaer and Vernhout,<br />

1902a, p. 3-13; Java—Lindgren, 1897, p. 485; 1898, p. 45; Ternate, East Indies—Thiele,<br />

1900, p. 72 (as Placospongia mixta); Torres Strait, between Australia and New Guinea<br />

—Ridley, 1884, p. 481; Aru Is. near New Guinea—Hentschel, 1912, p. 324. Tropical<br />

Pacific America: Punta Arenas, Costa Rica—Carter, 1880, p. 53 (as Placospongia sp.);<br />

Sollas, 1888, p. 272 (as Placospongia intermedia); Pacific coast <strong>of</strong> Panama—de Laubenfels,<br />

1936b, p. 454 (as Placospongia intermedia).<br />

DISCUSSION. Bowerbank established his Geodia carinata for specimens from the South<br />

Seas with an ectochrote <strong>of</strong> multiangulated cylindrical spicules and endosomal spirasters.<br />

He figured only an extreme modification <strong>of</strong> the latter microsclere, noting that it was<br />

<strong>of</strong>ten more elongate in form. In both size and form, the spicules <strong>of</strong> the Jamaican sponges<br />

agree well with previous descriptions. Due to the contracted condition <strong>of</strong> my specimens,<br />

it was impossible to determine whether the smaller tylostyles were cortical.<br />

P. carinata has not been reported previously from the West Indies. Sollas did, however,<br />

describe a species, P. intermedia, from the Pacific Coast <strong>of</strong> Costa Rica. Unfortunately,<br />

he gave a confused description <strong>of</strong> the microscleres in his account <strong>of</strong> P. intermedia<br />

and in his redescription <strong>of</strong> P. carinata. As Lindgren (1898) has pointed out, Sollas almost<br />

certainly included the developmental stages <strong>of</strong> sterrasters under the heading <strong>of</strong> microstrongyles.<br />

Furthermore, after emphasizing the absence <strong>of</strong> spirasters in P. intermedia,<br />

he described spirastral microstrongyles, including some with large conical spines. Spirasters<br />

with such unbranched rays occur in my specimens and have been reported by<br />

Vosmaer and Vernhout for P. carinata. Carter (1880), in a discussion <strong>of</strong> P. melobesioides,<br />

mentions a species or variety with spirastral flesh spicules. The variety included only two<br />

specimens, one being the Geodia carinata <strong>of</strong> Bowerbank and the other a British <strong>Museum</strong><br />

specimen from Punta Arenas, Costa Rica. The second specimen is almost certainly the<br />

one on which Sollas* P. intermedia is based. To judge from Carter's description, multirayed<br />

spirasters occurred in both the Central American and South Seas examples.<br />

The only other distinguishing characteristics <strong>of</strong> P. intermedia Sollas are the presence<br />

<strong>of</strong> spherasters (a variable feature) and the small sizes (no ranges are given) <strong>of</strong> the tylostyles<br />

and the sterrasters as listed by Sollas. The Panamanian specimens attributed to<br />

P. intermedia by de Laubenfels also had small tylostyles and sterrasters. Their spirasters<br />

were said to have tylote or strongylote rays. De Laubenfels further reported, without<br />

comment, that the sterrasters were derived not from cylindrical spicules but from spiny<br />

spheres. Sollas had considered similar spheres to be the early stages <strong>of</strong> spherasters, which<br />

were, however, absent in the Panamanian specimens. De Laubenfels neither mentioned<br />

the presence nor noted the absence <strong>of</strong> an ectochrote. On the available evidence, P. intermedia<br />

Sollas is probably conspecific with P. carinata. The Jamaican sponges cannot be<br />

distinguished from the East Indian representative <strong>of</strong> P. carinata.<br />

Placospongia melobesioides Gray, another Indo-Pacific species, has been reported from<br />

the West Indies by Schmidt (1870), de Laubenfels (1936a) and Arndt (1927). Arndt's<br />

record is <strong>of</strong> a Curasao sponge with an ectochrote <strong>of</strong> spheres and an endosome devoid<br />

<strong>of</strong> spirasters. It seems clearly conspecific with P. melobesioides as redescribed by Carter<br />

(1880). De Laubenfels' minute specimen from the Dry Tortugas included nothing but<br />

sterrasters and must be considered unrecognizable. His identification was apparently<br />

made on the basis <strong>of</strong> Schmidt's dubious record from Florida. Schmidt's specimen is<br />

inadequately described. Sollas (1888) was unable to find spheres in Schmidt's type slides<br />

at the British <strong>Museum</strong>, but, as Arndt points out, the sample may not have included the<br />

outer cortex. A cylindrical specimen from Florida in the <strong>Yale</strong> <strong>Peabody</strong> <strong>Museum</strong>, identified<br />

as P. melobesioides by Schmidt, has the spiculation <strong>of</strong> P. carinata.


SYSTEMATICS 65<br />

ORDER EPIPOLASIDA Sollas 1888, sensu de Laubenfels, 1936a<br />

FAMILY TETHYIDAE Gray, sensu de Laubenfels, 1936a<br />

GENUS TETHYA Lamarck<br />

Tethya seychellensis (Wright) Sollas<br />

Alemo seychellensis Wright, 1881, p. 13 [type: Seychelles; Brit. Mus. (Nat. Hist. (?)].<br />

Tethya seychellensis, Sollas, 1888, p. 427; Keller, 1891a, p. 329; (?) Topsent, 1893, p. 177;<br />

Kirkpatrick, 1900a, p. 133; Wilson, 1902a, p. 388; Row, 1911, p. 303.<br />

Donatia seychellensis, Dendy, 1916a, p. 100; idem, 1916b, p. 265.<br />

Donatia ingalli var. seychellensis, Wilson, 1925, p. 335.<br />

Tethya ingalli, Lindgren (not Bowerbank, 1872), 1898, p. 36.<br />

Tethya maza, I. Sollas (not Selenka, 1880), 1902, p. 216.<br />

Tethya lyncurium var. C Dendy, 1905, p. 114 (fide Dendy, 1916b.)<br />

Donatia viridis Baer, 1906, p. 26.<br />

HABITAT. Common on turtle grass near Port Royal.<br />

SHAPE. Spherical to slightly ovoid. Budding specimens are 1.5-4 cm in diameter.<br />

COLOR. The living sponges are a dull orange or a dull red. In alcohol, the sponges<br />

are beige, with or without an orange tinge.<br />

CONSISTENCY. Tough, slightly compressible.<br />

SURFACE. The surface is raised into low, flattened to rounded tubercles which are<br />

1-3 mm in diameter. Spherical to elongate buds in various stages <strong>of</strong> separation are present<br />

on most specimens. Mature specimens, and also buds, <strong>of</strong>ten have long thin processes<br />

which in some cases are attached to the turtle grass. The processes are usually less than<br />

0.5 mm in diameter but are <strong>of</strong>ten greater than 1 cm in length. A few specimens have<br />

small apical oscules, 1-2 mm in diameter, but most have no obvious exhalant openings.<br />

A network <strong>of</strong> narrow grooves runs between the tubercles. On their surface is a reticulation<br />

<strong>of</strong> bands visible to the unaided eye.<br />

ECTOSOME. The four-layered cortex includes an ectochrote, a cavernous layer, a spherastral<br />

layer, and a basal fibrous layer. The cortex is <strong>of</strong>ten more than a millimeter in total<br />

thickness. The tubercles are covered by the ectochrote and supported by gradually<br />

expanding spicule bundles. The ectochrote, which also covers the grooves, is composed<br />

<strong>of</strong> closely packed tylasters. The surface bands are 30-100 p in width. The interstices <strong>of</strong> the<br />

network, (100-<strong>20</strong>0 p in width) are pierced by numerous ostia, 30-45 p in diameter. At the<br />

grooves, the ectochrote overlies a cavernous layer in which the extensive cortical cavities<br />

are 450-1725 p in height. Vertical canals, 15-60 //, in diameter, run from the cavities to<br />

the endosome. The flesh in the cavernous layer contains sparsely scattered tylasters. A<br />

few spherasters may be present near the spicule bundles. The deeper part <strong>of</strong> the cortex<br />

contains tylasters and numerous spherasters. Microscleres are particularly abundant in the<br />

walls <strong>of</strong> the vertical canals. A basal fibrous layer, 30-70 p in thickness, contains sparsely<br />

scattered microscleres.<br />

ENDOSOME. Spicule bundles and individual megascleres are radially arranged around<br />

the central focal region. The latter, a confused mass <strong>of</strong> megascleres, is several millimeters<br />

in diameter. The endosomal microsclere complement includes spherasters, tylasters and<br />

oxyasters. The first category is rare and occurs only at the periphery. Tylasters become<br />

infrequent toward the center. Oxyasters are abundant in the flesh between spicule<br />

bundles. Attached buds are similar in structure to the parents but contain fewer oxyasters.<br />

The endosome <strong>of</strong> a specimen collected on January 5, 1961, contains closely<br />

packed, slightly ovoid eggs, 60-70 p in longest diameter. Their nuclei are about 13 JX in<br />

diameter.<br />

SPICULES, (a) Styles, ranging to strongyloxeas, straight, 328-1788 X 5-35 p. The shaft<br />

has its maximum diameter in mid-length. The apices <strong>of</strong> stylote spicules are <strong>of</strong>ten eroded<br />

in appearance, (b) Spherasters, with prominent conical rays nearly equal in length to<br />

the centrum diameter, 27-76 p, in overall diameter. The rays may be sharp, blunt or<br />

bifid, (c) Oxyasters, with 6-8 thin roughened rays, 24-83 p in total diameter. The small


66 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

central region is about 3 JU in diameter. The rays are straight, curved, somewhat irregular<br />

or bent. They have a roughened to spinous surface. In a few spicules, one or more<br />

rays are branched either terminally or at mid-length, (d) Tylasters, with 6-10 short,<br />

straight rays, 10-16 ^ in total diameter. The central region is 2 /x in diameter. The rays<br />

end in prominent roughened knobs.<br />

Individual analyses, lengths in microns:<br />

Strongyloxeas and Styles<br />

Spherasters<br />

328-1788 (50)<br />

27-73 (25)<br />

410-1568 (50)<br />

35-76 (25)<br />

500-1232 (25)<br />

35-72 (50)<br />

Tylasters<br />

Oxyaslers<br />

10-16 (25)<br />

45-83 (25)<br />

10-13 (25)<br />

29-37 (25)<br />

10-13 (25)<br />

24-37 (25)<br />

DISTRIBUTION. Tropical Atlantic America: Puerto Rico—Wilson, 1902a, p. 388. Indo-<br />

Pacific: Red Sea—Keller, 1891a, p. 329; Row, 1911, p. 303; Indian Ocean—Dendy, 1916b,<br />

p. 265; Christmas Is.—Kirkpatrick, 1900a, p. 133; Seychelles—Wright, 1881, p. 13 (as<br />

Alemo seychellensis); Northwest India—Dendy, 1916a, p. 100; Ceylon—Dendy, 1905, p.<br />

114; Malaya—I. Sollas, 1902, p. 216 (as Tethya maza); East Indies—Sollas, 1888, p. 427;<br />

Lindgren, 1898, p. 36 (as Tethya ingalli); Philippines—Sollas, 1888, p. 427; Wilson, 1925,<br />

p. 335; Tahiti, Pacific Ocean—Baer, 1906, p. 26 (as Donatia viridis).<br />

DISCUSSION. Despite several revisions, the taxonomy <strong>of</strong> Tethya remains in a chaotic<br />

state. The range <strong>of</strong> intraspecific variation remains uncertain, particularly with respect<br />

to cortical structure. Most records <strong>of</strong> oxyaster-containing sponges are difficult to evaluate,<br />

from lack <strong>of</strong> data. Many specimens, briefly described, have been attributed to poorly<br />

known species. The published synonymies are conflicting and <strong>of</strong>ten based on spicule<br />

data alone. The Jamaican sponges are very similar in spiculation and cortical structure<br />

to T. seychellensis (Wright), as described by Sollas in the Challenger Report. It should<br />

be noted that Wright's original description is not detailed and that his type material<br />

has been lost. The taxonomic summary includes only those records which indicate the<br />

presence <strong>of</strong> a definite lacunar layer in the cortex. Tethya ingalli Bowerbank (1872) and<br />

T. actinia de Laubenfels (1950a) have endosomal oxyasters as in T. seychellensis. Both<br />

species differ from Wright's and from the present specimens in having a completely<br />

fibrous cortex.<br />

Topsent (1918) and Burton (1924a) have attributed oxyaster-containing specimens <strong>of</strong><br />

Tethya to T. diploderma Schmidt <strong>of</strong> the West Indies. Schmidt's original description<br />

does not give a detailed account <strong>of</strong> cortical structure. He makes no mention <strong>of</strong> oxyasters.<br />

Topsent (1918, p. 574) assumed that oxyasters were present in Schmidt's material. Burton<br />

(1924a) claimed to have examined the types <strong>of</strong> T. diploderma Schmidt and T. seychellensis<br />

(Wright) in his revision. Wright, however, clearly stated (1881, p. 19) that his type<br />

material had been lost before the publication <strong>of</strong> his paper. Assuming that Schmidt's type<br />

material is in the British <strong>Museum</strong> Collection, no published evidence is available on the<br />

cortical structure <strong>of</strong> his species. (Burton included both T. ingalli, with a fibrous cortex,<br />

and T. seychellensis, with a multi-layered cortex, in the synonymy <strong>of</strong> T. diploderma.)<br />

At least two oxyaster-containing species <strong>of</strong> Tethya (T. actinia and T. seychellensis) occur<br />

in the region studied by Schmidt. Furthermore, sponges without oxyasters occur in the<br />

same area. (cf. Tethya seychellensis <strong>of</strong> de Laubenfels 1936a, p. 164). It is not impossible<br />

that Schmidt's original description <strong>of</strong> the microsclere complement is accurate. At present,<br />

T. diploderma Schmidt must be considered as unrecognizable.<br />

Tethya actinia de Laubenfels<br />

Tethya actinia de Laubenfels, 1950a, p. 116 [type: Bermuda; Brit. Mus. (Nat. Hist.) reg.<br />

no. 1948.8.6.48]; idem, 1954a, p. 234 (?).<br />

Tethya diploderma, de Laubenfels (not Schmidt, 1870), 1953a, p. 545 (?).


SYSTEMATICS 67<br />

HABITAT. The single specimen was collected in a few feet <strong>of</strong> water at Drunkenman's<br />

Cay. Unfortunately, only a small fragment was kept for sectioning.<br />

SHAPE. Small, spherical, about 1 cm in diameter.<br />

COLOR. The sponge had a greenish cortex, presumably due to algae, and an orange<br />

endosome.<br />

CONSISTENCY. Tough.<br />

SURFACE. The surface is covered with low, flat tubercles which are about 1 mm in<br />

diameter.<br />

ECTOSOME. An ectochrote <strong>of</strong> tylasters covers the entire surface. The tubercles are supported<br />

by gradually expanding spicule bundles. Some megascleres project beyond the<br />

ectochrote. The cortex is fibrous between the bundles with the fibrous tissue most conspicuous<br />

in the deeper half. Tylasters are abundant throughout the cortex. Spherasters<br />

are present in the peripheral part <strong>of</strong> the cortex but are uncommon in the deeper half.<br />

Most, if not all, <strong>of</strong> the spherasters in the latter region are associated with spicule tracts<br />

or cortical canals. A few cortical cavities occur in the middle and basal regions <strong>of</strong> the<br />

cortex. Narrow vertical canals run down to the cavities from the ectochrote, while others<br />

lead from the cavities to the endosome. Other canals (excurrent?) run vertically across<br />

the entire cortex.<br />

ENDOSOME. The periphery <strong>of</strong> the endosome contains numerous microscleres <strong>of</strong> all<br />

types. In the interior, the flesh contains many oxyasters and less numerous tylasters.<br />

Spicule bundles and individual megascleres radiate outward from the focal area.<br />

SPICULES, (a) Strongyloxeas, sometimes actually styles, straight, usually with unequal<br />

ends, 529-1478 x 10-27 ju (55 spicules), (b) Spherasters, typically with sharp, conical,<br />

simple rays, 29-50 ^ in total diameter (25 spicules). The rays are <strong>of</strong>ten nearly equal in<br />

length to the diameter <strong>of</strong> the centrum, (c) Tylasters, with 6-8 knobbed rays, with or<br />

without a small centrum, 9-13 fx in total diameter (25 spicules), (d) Oxyasters, usually<br />

with 6 slender, pointed rays, 34-61 ^ in total diameter (25 spicules). The small centrum<br />

is 2-5 ju, in diameter. The rays may be straight, curved, or bent. Most spicules have one<br />

forked ray, and some have two or three such rays. An occasional ray has a sub terminal<br />

branch, sometimes in addition to a terminal division. The rays are roughened in at least<br />

some oxyasters. A few spicules have blunt rather than pointed ends.<br />

DISTRIBUTION. Tropical Atlantic America: Bermuda—de Laubenfels, 1950a, p. 116;<br />

Gulf <strong>of</strong> Mexico—de Laubenfels, 1953a, p. 545 (as Tethya diploderma)}; Indo-Pacific—<br />

de Laubenfels, 1954a, p. 234 (?).<br />

DISCUSSION. The single specimen is similar in appearance to the common cay species<br />

and was unfortunately mistaken for it. The specimen is similar in structure and spiculation<br />

to Bermudan specimens in the <strong>Yale</strong> <strong>Peabody</strong> <strong>Museum</strong> identified as T. actinia by<br />

de Laubenfels. The <strong>Peabody</strong> <strong>Museum</strong> specimens have knobbed tylasters. De Laubenfels<br />

(1950a, fig. 52) figured the small microscleres as strongylasters. The spherasters <strong>of</strong> the<br />

Jamaican sponge have a somewhat greater maximum diameter. The cortical structure<br />

<strong>of</strong> de Laubenfels* Pacific sponge was not described in detail.<br />

Tethya actinia is very similar to T. ingalli Bowerbank. An extensive geographical<br />

range has been attributed to the latter species, which was originally described from<br />

Australia. T. actinia differs from T. ingalli in having smaller spherasters and more consistently<br />

forked oxyasters. T. seychellensis (Wright), which <strong>of</strong>ten contains forked oxyasters,<br />

differs markedly from T. actinia in cortical structure.<br />

Tethya sp., cf. maza Selenka<br />

HABITAT. Common on the underside <strong>of</strong> rocks at the cays.<br />

SHAPE. The small hemispherical sponges are attached to the substratum by a flattened<br />

base. Typical specimens are 0.5-1 cm in diameter. Budding was not observed.<br />

COLOR. The living sponges are a pale yellow or pale orange. They become cream to<br />

<strong>of</strong>f-white in alcohol.<br />

CONSISTENCY. Tough, very slightly compressible.<br />

SURFACE. Much <strong>of</strong> the surface is covered by very low tubercles, 0.5-1.5 mm in<br />

diameter. No large oscules were seen.


68 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

ECTOSOME. The cortex is usually 0.5-1 mm in thickness. The ectochrote, 15-30 p in<br />

thickness, is packed with strongylasters. The tubercles are supported by the gradually<br />

expanding ends <strong>of</strong> choanosomal spicule bundles. The cortical megascleres are similar in<br />

size range to those <strong>of</strong> the interior. A few megascleres project beyond the ectochrote.<br />

Between the spicule bundles, the cortex contains scattered strongylasters and an abundance<br />

<strong>of</strong> spherasters. The cortex is not conspicuously fibrous. Cortical cavities are<br />

occasionally present but are not localized at a particular level.<br />

ENDOSOME. Stout spicule bundles, <strong>20</strong>0-300 ^ in diameter, radiate outward from a<br />

confused focal area. Individual radially-oriented megascleres are also present. A few<br />

spherasters are present, particularly at the periphery. Strongylasters are scattered throughout<br />

the interior and are particularly abundant near the canals.<br />

SPICULES, (a) The megascleres are mostly strongyloxeas. A few styles are present, 304-<br />

1318 x 5-22 fi. (b) Spherasters, with conical sharp-pointed rays, 15-82 fi in total diameter.<br />

An occasional ray is blunt or bifid. The ray length is less than the centrum diameter.<br />

(c) Strongylasters, with 8-12 rays, with or without a globular centrum, 10-17 fi in total<br />

diameter. The thin cylindrical rays are typically 5 p long and less than 2 ^ in diameter.<br />

They are sometimes slightly tylote and are rarely forked. In one <strong>of</strong> 4 analyzed specimens,<br />

the endosomal micrasters have a slightly larger maximum diameter (17 fx) than the<br />

ectosomal ones. (15 ju,).<br />

Individual analyses, lengths in microns:<br />

Strongyloxeas Spherasters Strongylasters<br />

304-1318 (50) 36-72 (35) 10-15 (50)<br />

433-1240 (25) 35-82 (50) 12-13 (50)<br />

present 22-58 (25) 12-17 (25)<br />

present 15-58 (25) 10-13 (25)<br />

DISCUSSION. If the original description is accurate, the Jamaican sponges are conspecific<br />

with Tethya maza Selenka (1879, p. 472) <strong>of</strong> Brazil.<br />

Sollas (1888, p. 440) reported finding long-rayed micrasters in "typical" specimens <strong>of</strong><br />

T. maza obtained from E. P. Wright. Wright presumably received the sponges from<br />

Selenka, with whom he was in communication. Burton (1924a, p. 1040) apparently<br />

considered the "typical" material as including the type specimen. He retained T. maza<br />

as a valid species, characterized by the frequent reduction <strong>of</strong> rays in many microscleres<br />

(cf. Burton, 1924a, plate 1, fig. 3).<br />

The Jamaican sponges are clearly not conspecific with those <strong>of</strong> Burton. It is by no<br />

means certain, however, that the British <strong>Museum</strong> collection includes the type specimen.<br />

The similarities between the Jamaican sponges and the original description <strong>of</strong> T. maza<br />

suggest that the specimens available to Sollas and Burton are representatives <strong>of</strong> a second<br />

Brazilian species.<br />

ORDER CHORISTIDA Sollas, 1888, sensu LeVi, 1956b<br />

FAMILY GEODIIDAE Gray, sensu de Laubenfels, 1936a<br />

GENUS GEODIA Lamarck<br />

SUBGENUS Geodia Lamarck<br />

The incurrent chones have cribriporal ro<strong>of</strong>s. The conspicuous oscules are localized<br />

in clusters.<br />

Geodia (Geodia) gibberosa Lamarck<br />

(Plate VIII, fig. 2)<br />

Geodia gibberosa Lamarck, 1815, p. 324 [type: Guiana; Mus. nat. Hist. nat. Paris];<br />

Duchassaing and Michelotti, 1864, p. 105; Bowerbank, 1873, p. 6; Carter, 1882, p.<br />

362; Topsent, 1889, p. 34, 48; idem, 1918, p. 612; idem, 1930, p. 3; Arndt, 1927, p.


SYSTEMATICS 69<br />

137; de Laubenfels, 1936a, p. 172; idem, 1936b, p. 454; idem, 1939, p. 6; idem, 1949,<br />

p. 27; idem, 1950a, p. 125; idem, 1953a, p. 551; idem, 1956, p. 2; LeVi, 1959, p. 117;<br />

Wells et al., 1960, p. 235; Little, 1963, p. 59.<br />

Pyxitis gibberosa, Schmidt, 1870, p. 70.<br />

Geodia cariboa [sic] Duchassaing and Michelotti, 1864, p. 105.<br />

Geodia tuberculosa Bowerbank, 1872, p. 626; Carter, 1884, p. <strong>20</strong>8; Sottas, 1888, p. 251.<br />

Geodia tumulosa Bowerbank, 1872, p. 628; Sollas, 1888, p. 249.<br />

Geodia media Bowerbank, 1873, p. 13; Uliczka, 1929, p. 56.<br />

Synops (?) media, Sollas, 1888, p. 266.<br />

Geodia reticulata Bowerbank, 1874a, p. 300; Sollas 1888, p. 253.<br />

HABITAT. The sponges are common on ro<strong>of</strong>s along the mangrove boat channels.<br />

Other specimens were dredged from Pigeon House Pile.<br />

SHAPE. Massive, <strong>of</strong>ten lobate. Many specimens are more than 10 cm in diameter. The<br />

broad lobes are usually several centimeters in height.<br />

COLOR. The surface varies in color between a near-white and a dark brown. Many<br />

individuals are mottled. The oscular areas are darker than the rest <strong>of</strong> the surface and<br />

may be nearly black. The cortex, at least in its deeper parts, is white. The interior is<br />

beige. The sponges undergo little change in alcohol.<br />

CONSISTENCY. The hard cortex gives the sponge a leathery to stony consistency. The<br />

interior is tough and compact.<br />

SURFACE. Even to uneven. The sponge has one to several oscular areas which are<br />

<strong>of</strong>ten apical on flattened lobes. The oscular areas are typically circular and 1-2 cm in<br />

diameter. They are occasionally ovoid, with the maximum diameter attaining 8 cm. The<br />

individual openings are usually less than 0.5 mm in diameter but occasionally reach a<br />

diameter <strong>of</strong> 1.5 mm.<br />

ECTOSOME. The ectosome is a thick three-layered cortex. Extensive areas <strong>of</strong> the surface<br />

are cribriporal due to the presence <strong>of</strong> numerous dermal pores. Individual ostia are<br />

<strong>20</strong>-50 fx in diameter. They are separated by narrow bands <strong>of</strong> flesh which contain numerous<br />

small strongylospherasters. Small oxeas are radially arranged in the outer cortex and<br />

project beyond the ectochrote. The sterrastral crust is crossed by incurrent chone canals<br />

which are 50-100 //, in diameter. Adjacent canals are typically 0.5 mm apart. Their fibrous<br />

linings contain numerous strongylospherasters. The canals become compressed by large<br />

sphincters in the fibrous layer <strong>of</strong> the cortex. Large subcortical spaces are present beneath<br />

the sphincters. Under the oscular areas, the sterrastral crust is crossed by exhalant<br />

canals which are 300-600 ^ in diameter. Endosomal megascleres occasionally project into<br />

the cortex.<br />

ENDOSOME. Cavernous. Individual megascleres and spicule bundles are radially arranged<br />

in the periphery <strong>of</strong> the endosome. The triaene cladomes are tangentially placed<br />

in the fibrous layer <strong>of</strong> the cortex. In the deeper parts <strong>of</strong> the interior, the megascleres<br />

and bundles are scattered in confusion. The flesh contains numerous oxyasters and all<br />

stages <strong>of</strong> sterrasters. Strongylospherasters are most abundant toward the periphery <strong>of</strong><br />

the endosome. Some, but not all <strong>of</strong> the subcortical strongylospherasters have a distinctive<br />

size and shape.<br />

SPICULES, (a) Plagiotriaenes, <strong>of</strong> greatly varied size, with rhabds 290-15<strong>20</strong> X 10-40 /A,<br />

and clads 21-304 X 12-35 /x. Protriaenes, anatriaenes, mesodiaenes and mesomonaenes<br />

occur sporadically and in low numbers. It is possible that some are not proper to the<br />

sponge, (b) Large oxeas, slightly curved, 714-1972 X 7-43 jU(. A few styles also occur.<br />

(c) Small cortical oxeas, thin, slightly curved, gradually pointed, 137-224 x 3-4 fi. (d)<br />

Sterrasters, spherical to slightly ovoid, 53-108 ^ in longest diameter. The surface <strong>of</strong><br />

mature spicules is covered by flattened, polygonal tubercles which are typically 3 /x in<br />

diameter. Spiny developmental sterrasters are abundant in the interior, (e) Strongylospherasters,<br />

with a centrum and 5-8 short blunt rays, 5-8 /x in overall diameter, (f) Some<br />

<strong>of</strong> the subcortical strongylospherasters, 9-12 //, in diameter, have a centrum and 10-12<br />

rays. Intermediates between subcortical and typical strongylospherasters occur, (g) Oxyasters,<br />

with 5-10 rays, with or without a small centrum, 15-33 fi in total diameter. The


70 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

rays are slender, cylindrical, and slightly roughened. Individual analyses, lengths in<br />

microns:<br />

Plagiotriaenes (rhabds)<br />

290-1378<br />

492-15<strong>20</strong><br />

550-1440<br />

714-1229<br />

Small oxeas<br />

159-210 (25)<br />

174-210 (25)<br />

166-224 (25)<br />

137-188 (25)<br />

Subcortical<br />

strongylospher asters<br />

present<br />

9-12 (25)<br />

9-12 (25)<br />

present<br />

(dads)<br />

21-232 (5)<br />

53-304 (10)<br />

117-258 (10)<br />

82-211 (10)<br />

Sterrasters<br />

77-108 (25)<br />

64-82 (50)<br />

53-82 (50)<br />

64-88 (25)<br />

Oxyasters<br />

15-<strong>20</strong> (25)<br />

16-26 (25)<br />

16-33 (25)<br />

16-30 (25)<br />

Large Oxeas<br />

1088-1972 (12)<br />

749-1345 (25)<br />

714-1800 (25)<br />

737-1312 (25)<br />

Strongylospherasters<br />

5-7 (25)<br />

5-8 (25)<br />

5-8 (25)<br />

5-8 (25)<br />

DISTRIBUTION. Tropical Atlantic America: North Carolina—Wells et al., 1960, p. 235;<br />

South Carolina—Schmidt, 1870, p. 70; Florida—Schmidt, 1870, p. 70 (both as Pyxitis<br />

gibberosa); Carter, 1884, p. <strong>20</strong>8 (as Geodia tuberculosa)) Dry Tortugas, Florida—de<br />

Laubenfels, 1936a, p. 172; West Coast, Florida—de Laubenfels, 1953a, p. 551 and Little,<br />

1963, p. 59; Gulf <strong>of</strong> Campeche, Mexico—Topsent, 1889, p. 34, 48; Mexico—Bowerbank,<br />

1872, p. 626 (as Geodia tuberculosa); Bowerbank, 1873, p. 13 (as Geodia media); Bowerbank,<br />

1874a, p. 300 (as Geodia reticulata); Honduras—Bowerbank, 1872, p. 628 (as<br />

Geodia tumulosa); Atlantic coast, Panama—de Laubenfels, 1936b, p. 454; Venezuela—<br />

Carter, 1882, p. 362; Guiana—Lamarck, 1815, p. 324 (?); Brazil—de Laubenfels, 1956, p.<br />

2; Bermuda—de Laubenfels, 1950a, p. 125; Bahamas—de Laubenfels, 1949, p. 22; Cuba<br />

—Schmidt, 1870, p. 70 (as Pyxitis gibberosa); Jamaica—Bowerbank, 1872, p. 628 (as<br />

Geodia tumultosa); Antilles—Schmidt, 1870, p. 70 (as Pyxitis gibberosa); Lesser Antilles:<br />

Tortuga—Uliczka, 1929, p. 56 (as Geodia media); Tortola, Br. Virgin Is.—Duchassaing<br />

and Michelotti, 1864, p. 105; St. Thomas—Duchassaing and Michelotti, 1864, p. 105<br />

(as Geodia cariboo); Gaudeloupe—Topsent, 1889, p. 34, 48; Dominica—Bowerbank,<br />

1873, p. 6; St. Vincent—Carter, 1882, p. 362; Barbados—Uliczka, 1929, p. 56 (as Geodia<br />

media); Curasao—Arndt, 1927, p. 137: Old Providence Is., Caribbean—de Laubenfels,<br />

1939, p. 6. Tropical West Africa: Sao Tome—Topsent, 1918, p. 612; LeVi, 1959, p. 117.<br />

Tropical Pacific America: Pacific coast, Panama—de Laubenfels, 1936b, p. 454.<br />

DISCUSSION. The presence <strong>of</strong> cortical oxeas and an ectochrote has been overlooked in<br />

recent studies. The Jamaican sponges agree well in spiculation with Lamarck's type which<br />

was restudied by Topsent (1918, 1930). Bowerbank, who also examined Lamarck's specimen<br />

(1873), described five new species <strong>of</strong> Geodia from Tropical Atlantic American<br />

waters (1873-75). His species are all very similar in structure and spiculation. Geodia<br />

dysoni Bowerbank, 1873, differs from the others and from G. gibberosa in having dispersed<br />

oscules. The remaining species are probably synonyms <strong>of</strong> G. gibberosa. A reexamination<br />

<strong>of</strong> type specimens would be desirable. Anatriaenes and protriaenes were<br />

reported for several <strong>of</strong> Bowerbank's species. Such spicule categories were occasionally<br />

present in the Jamaican material. Uliczka's Jamaican record <strong>of</strong> Geodia media Bowerbank<br />

certainly refers to Geodia gibberosa.<br />

GENUS GEODIA Lamarck<br />

SUBGENUS Cydonium Fleming<br />

Spicules as in the typical subgenus. The incurrent and excurrent openings are<br />

cribriporal and similar in size.


SYSTEMATICS 71<br />

Sollas (1888) used Cydonium as a genus for geodiids in which the oscules and pores<br />

could not be distinguished. Sollas' classification <strong>of</strong> the family Geodiidae merits reconsideration.<br />

Geodia (Cydonium) papyracea n. sp.<br />

(Text-fig. 13; plate VIII, figs. 1, 2)<br />

Fig. 13.—Spicules <strong>of</strong> Geodia papyracea. A. Oxeas. B. Distal end <strong>of</strong> plagiotriaene. C. Distal end<br />

<strong>of</strong> anatriaene. D. Small spherasters. E. Thick-rayed subcortical oxyaster. F. Oxyaster. G.<br />

Sterrasters. H. Surface sculpturing <strong>of</strong> portions <strong>of</strong> sterrasters enlarged. A: scale I. B, C:<br />

scale II. D, E, F: scale III. G: scale IV. H: scale V. YPM 5045. Holotype.<br />

HOLOTYPE. YPM 5045. Mangrove boat channel, July 13, 1961.<br />

HABITAT. Common on mangrove roots along the main boat channel.<br />

SHAPE. Massive, <strong>of</strong>ten fist-sized. A few specimens attain a diameter <strong>of</strong> 8-10 cm.<br />

COLOR. The living sponges are <strong>of</strong>f-white to light gray. Purple tinges are sometimes<br />

present. Preserved specimens have similar colors.<br />

CONSISTENCY. Firm to compressible. When pressure is exerted many specimens collapse.<br />

The cortex is rather brittle. The endosome is pulpy and easily crumbled.<br />

SURFACE. Rough to the touch. The surface may be even, wrinkled, or folded. No<br />

macroscopic oscules are present. Considerable areas <strong>of</strong> the surface have a punctiform<br />

appearance. The sponge is hispid as seen under a dissecting microscope at a magnification<br />

<strong>of</strong> 60 x.<br />

ECTOSOME. The cortex is 400-600 /x in thickness. A thin ectochrote <strong>of</strong> strongylospherasters<br />

is pierced by openings which are 35-50 p in diameter. The openings form the<br />

cribriporal ro<strong>of</strong>s <strong>of</strong> chone canals. A thin lacunar layer separates the ectochrote from<br />

the sterrastral crust. The cavities are crossed by broad columns which contain megascleres<br />

and a few strongylospherasters. A distinct basal fibrous layer is absent. Numerous thick


72 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

plagiotriaenes project through the entire cortex. Their cladomes are inserted into the<br />

ectochrote. Large oxeas and thin anatriaenes are also present. A few triaenes have an<br />

oblique or tangential position in the outer cortex.<br />

ENDOSOME. The peripheral region has a radiate architecture, while the interior is<br />

confused. Most <strong>of</strong> the oxeas end within the endosome. The plagiotriaenes only infrequently<br />

terminate below the cortex. Their heads usually begin in the periphery <strong>of</strong> the<br />

endosome and enter the cortex singly or in loose groups. The flesh contains numerous<br />

sterrasters in all stages <strong>of</strong> development. Oxyasters and occasional strongylasters are also<br />

present. Distinctive multi-rayed oxyasters are present in the periphery.<br />

SPICULES, (a) Robust plagiotriaenes, with rhabds 406-1058 x 5-24 /x, and clads 33-<br />

123 x 4-24 /x. (b) Slender anatriaenes, with rhabds 246-609 X 2-3 /x, and clads 7-36 X<br />

1-2 jit. (c) Oxeas, straight to slightly curved, gradually to abruptly pointed, 616-1183<br />

X 9-<strong>20</strong> p. (d) Sterrasters, mostly spherical, occasionally somewhat ovoid, 52-75 /x in<br />

diameter. The surface <strong>of</strong> mature spicules is covered by star-shaped tubercles about 3 /x<br />

in diameter. Smaller, spiny developmental forms <strong>of</strong> sterrasters are present in the flesh, (e)<br />

Strongylospherasters, with 8-10 blunt rays, 5-7 /x in total diameter. The centrum is<br />

about 2 /x in diameter, (f) Oxyasters, with 6-12 thin, roughened rays, with or without a<br />

small centrum, 24-48 /x in total diameter. The rays are 1-2 /x in diameter, (g) Some, but<br />

not all <strong>of</strong> the subcortical oxyasters are <strong>of</strong> a distinctive category, with numerous roughened<br />

or minutely spined rays, 2-3 /x in diameter. The spicules have an overall diameter <strong>of</strong><br />

21-36 /x. Individual analyses, lengths in microns:<br />

Plagiotriaenes<br />

(rhabds)<br />

406-956*<br />

449-674<br />

782-1058<br />

Oxeas<br />

638-1058 (30)*<br />

624^1137 (25)<br />

616-1183 (15)<br />

{clads)<br />

36-101 (15)*<br />

33-87 (5)<br />

65-123 (10)<br />

Sterrasters<br />

50-72 (50)*<br />

58-72 (50)<br />

52-75 (25)<br />

Anatriaenes<br />

{rhabds)<br />

297-470*<br />

246-609<br />

297-500<br />

[clads)<br />

7-36 (6)*<br />

9-29 (25)<br />

10-35 (10)<br />

Strongylospherasters<br />

5-7 (25)*<br />

5-7 (25)<br />

5-7 (25)<br />

Thick-rayed<br />

Oxyasters<br />

subcortical oxyasters<br />

29-46 (25)*<br />

21-30 (25)*<br />

24-46 (25)<br />

22-36 (25)<br />

24-48 (25)<br />

26-34 (5)<br />

A few long, thin promesodiaenes are present in the third specimen.<br />

DISCUSSION. The species is characterized by its peculiar consistency, thin cortex, small<br />

megascleres, small sterrasters and subectochrotal spaces. The only other representative<br />

<strong>of</strong> the subgenus Cydonium in tropical Atlantic American waters is Geodia glariosus<br />

(Sollas, 1886, p. 196) <strong>of</strong> Bahia, Brazil. G. glariosus has very large triaenes, special cortical<br />

oxeas, and small choanosomal oxyasters. Its ectochrote overlies a layer <strong>of</strong> coarse sand.<br />

GENUS ERYLUS Gray<br />

Erylus ministrongylus n. sp.<br />

(Text-fig. 14; plate VIII, fig. 3)<br />

HOLOTYPE. YPM 5046. Drunkenman's Cay, July 6, 1961.<br />

HABITAT. TWO specimens were collected under rocks in a few feet <strong>of</strong> water at<br />

Drunkenman's Cay.<br />

SHAPE. The holotype is a conical mass with a maximum height <strong>of</strong> 2.5 cm. The second<br />

specimen is an incrustation, 1-3 mm in thickness.<br />

COLOR. The incrusting specimen was <strong>of</strong>f-white in life. Most <strong>of</strong> one side <strong>of</strong> the massive<br />

example was dark brown. The remainder <strong>of</strong> its surface was white, as was the interior<br />

<strong>of</strong> both sponges. The colors are retained in alcohol.


SYSTEMATICS 73<br />

Fig. 14.—Spicules <strong>of</strong> Erylus ministrongylus. A. Strongyles. B. Aspidasters. C. Centrotylote oxeas.<br />

D. Few-rayed oxyasters. E. Many-rayed oxyaster. A, B: scale I. C, D, E: scale II.<br />

YPM 5046. Holotype.<br />

CONSISTENCY. Very slightly compressible.<br />

SURFACE. The larger specimen has long broad folds and troughs. Much <strong>of</strong> its surface<br />

is riddled with small (inhalant?) openings less than 0.5 mm in diameter. The sponge<br />

has one apical and two lateral oscules, each about 3 mm in diameter. The small openings<br />

are absent from some areas, including the regions around the oscules and the tops <strong>of</strong><br />

many folds. The incrusting specimen has small openings over the entire surface and<br />

no large oscules. Its surface is even.<br />

ECTOSOME. The cortex, 150-<strong>20</strong>0 LL in thickness, can be detached in flakes. The cortical<br />

armor is built <strong>of</strong> many layers <strong>of</strong> tangentially placed disk spicules (aspidasters). Centrotylote<br />

oxeas form a sparse covering over the armor and a dense feltwork around the<br />

small surface openings. The uniporal openings lead to canals which are 40-<strong>20</strong>0 LL in<br />

diameter. The canal linings contain numerous multi-rayed asters. Smaller numbers <strong>of</strong><br />

oxeas and few-rayed asters are also present in the linings.<br />

ENDOSOME. The interior is packed with two types <strong>of</strong> euasters. Centrotylote oxeas are<br />

common subcortically but become rare in the interior. Strongylote megascleres are scattered<br />

singly and also grouped into bundles which are 40-150 p in diameter. Many<br />

bundles are radially or obliquely oriented. The triaenes are subcortical with their<br />

cladomes spreading under the cortex. A few aspidasters are present in the endosome.<br />

SPICULES, (a) Orthotriaenes and dicho-orthotriaenes, with a short rhabd. The rhabds<br />

are 187-311 X 12-17 LL, and the clads 119-319 x 5-13 LL. In most spicules, rhabd length<br />

is slightly greater than clad length. A few spicules seen in section have clads longer<br />

than the radially placed rhabd. As examples, triaenes with unbroken rhabds 187, 238,<br />

238, and 278 LL in length have clad lengths <strong>of</strong> 136, 211, 244, and 178 LL respectively.<br />

The clads may be straight, undulating, terminally bent, or dichotomous. The deuteroclads<br />

are usually very short. For example, clads with a protoclad length <strong>of</strong> 260, 196 and 140 LL<br />

have deuteroclads <strong>20</strong>, 10 and <strong>20</strong> LL long respectively, (b) Strongyles, straight to very<br />

slightly curved, 370-551 x 5-23 LL. The majority have rounded, but narrow ends. A few<br />

are typical blunt strongyles. A very few stylote and oxeote spicules occur, (c) Aspidasters


74 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

(disk-spicules). Thin, elliptical, 87 X 43 to 181 X 87 /x. The spicules are only 7-14 //, in<br />

thickness. Typical sizes are 137 X 79, 145 x 72, and 166 x 79 //,. Their outline is<br />

usually irregular but seldom jagged. The minute, thickly-strewn tubercles give the surface<br />

a granular appearance, (d) Smooth centrotylote oxeas, nearly straight to slightly<br />

curved, 37-80 X 3-5 jx. The central inflation is elongate and ill-defined. For example,<br />

a spicule with a mid-length diameter <strong>of</strong> 4 ^ has a diameter <strong>of</strong> 3 fi at a point adjacent<br />

to the swelling. A few spicules have lateral as well as mid-length swellings. The ends<br />

are almost always sharp-pointed. A single spicule found in the incrusting specimen had<br />

a roughened surface, (e) Few-rayed oxyasters, with 3-6 rays. Ray length varies between<br />

9 and 33 /x. The rays have a basal diameter <strong>of</strong> 2-3 p. They are straight, pointed and<br />

at least distally roughened. A few spicules have very thin rays which seem to be smooth.<br />

The central meeting point <strong>of</strong> the rays is only 3 //, in diameter, (f) Multi-rayed oxyasters,<br />

10-25 p in overall diameter. About a dozen rays project from a central region which is<br />

2-3 fM in diameter. The rays have a basal diameter <strong>of</strong> 1 /x. The larger spicules at least<br />

are minutely roughened. Individual analyses, lengths in microns: (holotype listed first<br />

in each category).<br />

Strongyloxeas Aspidasters Centrotylote oxeas<br />

398-534 (50)* 106 X 70-158 X 64 (25)* 53-80 (50)*<br />

370-551 (50) 87 X 43-181 X 87 (50) 37-61 (50)<br />

Few-rayed oxyasters (ray length) Multi-rayed oxyasters (total diam.)<br />

13-33 (25)* 10-25 (25)*<br />

9-26 (50) 12-24 (50)<br />

Triaene clad length Rhabd length<br />

136-260 (30)* 187-278 (5)*<br />

119-319 (25) 217-311 (5)<br />

DISCUSSION. The monaxonid megascleres are unusually small for the genus. The<br />

Jamaican sponge is related to Erylus euastrum (Schmidt, 1868) which, together with<br />

its apparent synonym, E. transiens (Weltner), has been reported from the West Indies.<br />

The Jamaican species differs from Schmidt's (as described by Babi£, 1922, p. 287 and<br />

Topsent, 1928, p. 115) in having smaller monaxonid spicules and roughened rather<br />

than smooth-rayed large asters. E. alleni de Laubenfels 1934, <strong>of</strong> Puerto Rico has smaller<br />

aspidasters, oxeote megascleres, and very small multi-rayed oxyasters. Its large asters<br />

have smooth rays.<br />

FAMILY CHONDRILLIDAE Gray, 1867<br />

GENUS CHONDRILLA Schmidt<br />

Chondrilla nucula Schmidt<br />

Chondrilla nucula Schmidt, 1862, p. 39 [type: Quarnero, Yugoslavia; repository unknown];<br />

idem, 1870, p. 26; Schulze, 1877a, p. 108; Carter, 1881, p. 249; idem, 1882,<br />

p. 268; idem, 1885, p. <strong>20</strong>0; idem, 1886a, p. 277; idem, 1890, p. 565; Topsent, 1890a,<br />

p. 27; idem, 1892, p. 56; idem, 1918, p. 601; idem, 1925a, p. 630; Keller, 1891a, p.<br />

327; Lendenfeld, 1897, p. 37; Wilson, 1902a, p. 386; Verrill, 1907, p. 334; Annandale,<br />

1915, p. 470; Babic, 1922, p. 269; Burton, 1924b, p. <strong>20</strong>6; idem, 1934, p. 522; idem,<br />

1936a, p. 7; Dendy, 1924, p. 314; Arndt, 1927, p. 138; Vosmaer, 1933, p. 293; de<br />

Laubenfels, 1936b, p. 464; idem, 1949, p. 22: idem, 1950a; p. 133; idem, 1954a, p. 250<br />

(?); idem, 1956, p. 2, 4; Sara, 1958a, p. 216; Sara and Siribelli, 1960, p. 31; LeVi,<br />

1959, p. 121; Vacelet, 1959, p. 46; idem, 1961, p. 30; Little, 1963, p. 63.<br />

[non] Chondrilla nucula, Kieschnick, 1896, p. 534 (fide Thiele, 1900, p. 63); de Laubenfels,<br />

1935a, p. 12.<br />

Chondrilla embolophora Schmidt, 1862, p. 39 (fide Topsent, 1918.)<br />

Chondrilla australiensis, de Laubenfels, 1954a, p. 249 (?) (not Carter, 1873.)


SYSTEMATICS 75<br />

HABITAT. Common on pilings and rocks at Port Royal and at the cays.<br />

SHAPE. The sponge spreads horizontally over the substratum, attaining a thickness<br />

<strong>of</strong> 1-2 mm.<br />

COLOR. A mottled brown to yellowish-brown, with pale areas around the oscules.<br />

The interior is drab. In alcohol, the original coloration is retained to a certain extent.<br />

CONSISTENCY. Tough.<br />

SURFACE. Smooth, shiny. The oscules are small, scattered and contractile.<br />

ECTOSOME. A detachable, densely pigmented skin is present. Spherasters <strong>of</strong> all sizes<br />

are scattered in it. Dermal pores could not be detected.<br />

ENDOSOME. The dense flesh is traversed by long, narrow, largely vertical canals, 10-25<br />

H in diameter. Microscleres are abundant only in the periphery <strong>of</strong> the endosome and<br />

in the vicinity <strong>of</strong> the large canals.<br />

SPICULES. Spherasters, typically with short, conical rays and with a definite centrum,<br />

10-40 JJL in total diameter. A varied number <strong>of</strong> spicules have a relatively small centrum<br />

and long oxeote rays which are as much as 12 /A in length. Spicules with short blunt<br />

rays also occur, again in varied numbers. In some specimens ray length as a percentage<br />

<strong>of</strong> centrum diameter seldom exceeds 25 per cent. In other specimens ray length is <strong>of</strong>ten<br />

equal to 40-60 per cent <strong>of</strong> centrum diameter and may even be greater than the centrum<br />

diameter. Individual analyses, total diameter in microns: 10-34 (75); 15-37 (50); <strong>20</strong>-40<br />

(50); 12-33 (50); <strong>20</strong>-40 (50).<br />

DISTRIBUTION. Tropical Atlantic America: Florida—Schmidt, 1870, p. 26; West Coast,<br />

Florida—Little, 1963, p. 63; Atlantic coast <strong>of</strong> Panama—de Laubenfels, 1936b, p. 464;<br />

Venezuela—Carter, 1882, p. 268; Brazil—de Laubenfels, 1956, p. 2, 4; Bermuda—<br />

Verrill, 1907, p. 334; de Laubenfels—1950a, p. 133; Bahamas—de Laubenfels, 1949, p.<br />

22; West Indies—Carter, 1881, p. 249; 1882, p. 268; Puerto Rico—Wilson, 1902a, p. 386;<br />

Antilles—Schmidt, 1870, p. 26; Curasao—Arndt, 1927, p. 138; Fernando Noronha,<br />

South Atlantic <strong>of</strong>f Brazil—Carter, 1890, p. 565; South Trinidad Is., South Atlantic, <strong>of</strong>f<br />

Brazil—Dendy, 1924, p. 314.<br />

Mediterranean-Atlantic: French Mediterranean—Vacelet, 1959, p. 46-48; Ligurean<br />

Sea, Italy—Sara, 1958a, p. 216; Naples, Italy—Topsent, 1925a, p. 630; Vosmaer, 1933,<br />

p. 293; Sara and Siribelli, 1960, p. 31; Adriatic—Schmidt, 1862, p. 39; 1862, p. 39<br />

(as Chondrilla embolophora); Schulze, 1877a, p. 108; Lendenfeld, 1897, p. 37; Babic\<br />

1922, p. 269; Mediterranean coast <strong>of</strong> Egypt—Burton, 1936a, p. 7; Corsica, Vacelet, 1961,<br />

p. 30; Azores—Topsent, 1890a, p. 27, 1892, p. 56; Gulf <strong>of</strong> Guinea—L^vi, 1959, p. 121;<br />

Sao Tom£, West Africa—Topsent, 1918, p. 601; Indo-Pacific: Red Sea—Carter, 1881,<br />

p. 249; Keller, 1891a, p. 327; Mauritus—Carter, 1881, p. 249; Gulf <strong>of</strong> Manaar, between<br />

India and Ceylon—Carter, 1881, p. 249; Mergui Archipelago, <strong>of</strong>f Burma—Annandale,<br />

1915, p. 470; Moluccas, East Indies—Carter, 1881, p. 249; Australia—Carter, 1885, p.<br />

<strong>20</strong>0, 1886a, p. 227; Great Barrier Reef, Australia—Burton, 1934, p. 522; West Central<br />

Pacific—de Laubenfels, 1954a, p. 250 (?); 1954a, p. 249 (as Chondrilla australiensis)}.<br />

DISCUSSION. Long-rayed spherasters occur in <strong>Yale</strong> <strong>Peabody</strong> <strong>Museum</strong> specimens <strong>of</strong><br />

Chondrilla nucula from Bermuda and the Virgin Islands. Schmidt (1870) reported them<br />

for his specimens from the Antilles and Florida. De Laubenfels (1935a) referred an<br />

aspiculous sponge from lower California to Chondrilla nucula. It was presumably a<br />

Chondrosia. De Laubenfels (1954a) recorded black Pacific sponges as C. nucula Schmidt<br />

and C. australiensis Carter (1873a). The latter identification is certainly incorrect, as<br />

the characteristic spined oxyasters <strong>of</strong> C. australiensis are absent. The long-rayed spicules<br />

<strong>of</strong> de Laubenfels' australiensis sponges are similar to those found in the above-mentioned<br />

West Indian specimens <strong>of</strong> C. nucula. It therefore seems probable that all <strong>of</strong> his black<br />

sponges are conspecific. As his sponges differ sharply from typical specimens <strong>of</strong> C. nucula<br />

in color (although not in spicule size), it is possible that they are a distinct species.<br />

ORDER HOMOSCLEROPHORA Levi 1953<br />

The Homosclerophora were dispersed in the classification <strong>of</strong> de Laubenfels (1936a).<br />

LeVi (1953, 1956b), in reconstituting the group, has emphasized its morphological and<br />

embryological uniformity.


76 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

FAMILY PLAKINIDAE Schulze, sensu Lendenfeld, 1903<br />

The families Plakinidae and Corticiidae as used by Sollas (1888) and LeVi (1953)<br />

are very similar and are here considered as a single group.<br />

GENUS PLAKORTIS Schulze<br />

Plakortis zyggompha (de Laubenfels) new combination<br />

Roosa zyggompha de Laubenfels, 1934, p. 2 [type: <strong>of</strong>f Puerto Rico; USNM No. 22277];<br />

idem, 1936a, p. 178.<br />

Plakortis simplex, de Laubenfels (not Schulze, 1880), 1950a, p. 132.<br />

HABITAT. Under rocks in shallow water at Maiden Cay.<br />

SHAPE. The seven specimens were small crusts, reaching a maximum thickness <strong>of</strong><br />

4.5 mm.<br />

COLOR. The exterior, in life and as preserved, is dark brown to drab. The interior<br />

is <strong>of</strong> a lighter shade than the surface.<br />

CONSISTENCY. Tough, rubbery, very slightly compressible.<br />

SURFACE. Smooth to the touch, even but microtuberculate as seen under a dissecting<br />

microscope. Five specimens have no obvious oscular openings. Each <strong>of</strong> the other two has<br />

a single small oscule.<br />

ECTOSOME. NO detachable dermis and little dermal specialization. The flesh becomes<br />

more compact towards the surface. A few possible dermal pores, <strong>20</strong>-60 /x in diameter, were<br />

observed in tangential sections taken at the surface. Narrow inhalant canals run downward<br />

toward the interior.<br />

ENDOSOME. The peripheral inhalant canals are collected by larger ones. The compact<br />

flesh is thickly strewn with spicules. Large convoluted embryos (proper to the sponge?)<br />

are present in specimens collected in July and August, 1961.<br />

SPICULES, (a) Diactinal spicules, with gradually pointed ends, 44-126 jx, in length and<br />

2-3 n in midlength diameter. The spicule is divided into two rays by a swollen, twisted,<br />

notched or lumpy region. The rays are almost always at an angle, giving the spicule a<br />

curved or sharply bent outline. The rays themselves may be bent. Thinner, developmental<br />

stages may be present. Individual analyses, length in microns, 75 diactinal spicules<br />

each: 56-115; 54-126; 63-117; 44-119. (b) Triactinal spicules, always less frequent<br />

than the diactinal ones, and rare in some specimens. The triactinals are sometimes regular<br />

and equiangular, but are more frequently sagittal and inequiangular in some degree.<br />

The rays are sharp-pointed, straight to curved and <strong>of</strong>ten twisted or lumpy at the base.<br />

A lumpy midregion occasionally has one or more short projections in addition to the<br />

usual 3 rays. Normal rays are 24-46 x 2-3 /x. (c) Rare Y-shaped and roughened ovoid<br />

spicules, 5-10 ju, in length. A few <strong>of</strong> the roughened spicules have short rays, indicating<br />

a derivation from the diactinal spicules.<br />

DISTRIBUTION. Tropical Atlantic America: Dry Tortugas, Florida—de Laubenfels,<br />

1936a, p. 178 (as Roosa zyggompha); Bermuda—de Laubenfels, 1950a, p. 132 (as Plakortis<br />

simplex) ?; Puerto Rico—de Laubenfels, 1934, p. 2 (as Roosa zyggompha).<br />

DISCUSSION. De Laubenfels (1934, p. 2) established Roosa zyggompha for Puerto Rican<br />

sponges similar in spiculation to Plakortis simplex Schulze. He later concluded (1950a, p.<br />

132) that the West Indian sponges were conspecific with Schulze's species. W. D. Hartman<br />

(personal communication) has pointed out the presence <strong>of</strong> minute spicules in West Indian<br />

Plakortis-like sponges. Such spicules are present, although rare, in U. S. National Muesum<br />

specimen number 22490. The specimen was collected at the Dry Tortugas and identified<br />

as Roosa zyggompha by de Laubenfels. The West Indian sponge is unlikely to be conspecific<br />

with Plakortis simplex Schulze. If the small microscleres are consistently present,<br />

and not developmental stages, Roosa de Laubenfels could be treated as a distinct genus.


DISTRIBUTION BY HABITAT 77<br />

GENUS CORTICIUM Schmidt<br />

Corticium tetralophum n. sp.<br />

(Text-fig. 15)<br />

HOLOTYPE. YPM 5047. Maiden Cay, Aug. 12, 1961.<br />

HABITAT. The single specimen was collected at Maiden Cay.<br />

SHAPE. A small incrustation with a maximum thickness <strong>of</strong> 2.5 mm.<br />

COLOR. The sponge was a dull yellow in life and is a grayish-beige in alcohol.<br />

CONSISTENCY. Compressible.<br />

SURFACE. The thin detachable ectosome contains numerous tetraloph spicules. A few<br />

triradiates are also present. No ostia could be found.<br />

ENDOSOME. Tetralophs and a few triradiates are sparsely scattered in the flesh. The<br />

ovoid flagellated chambers are 35-50 p in maximum diameter. Convoluted embryos<br />

(proper to the sponge?) are present in the specimen collected in early August.<br />

I I<br />

10//<br />

Fig. 15.—Spicules <strong>of</strong> Corticium tetralophum. Tetralophs. YPM 5047. Holotype.<br />

SPICULES, (a) Tetralophs, 17-30 //, in total diameter (100 spicules). The protoclads<br />

are 3 ^ in diameter. They bear 2-6, typically 4, deuteroclads. The latter are about equal<br />

in length to the protoclads. The smooth protoclads are 1-2 fi in basal diameter. Smooth<br />

and rough or spiny deuteroclads may be present on the same spicule. A typical spicule<br />

has 4 similar deuteroclads on each <strong>of</strong> the 4 rays. A rhabd cannot be distinguished from<br />

3 clads. (b) Very rare trilophate spicules, with the unbranched ray shortened or bent.<br />

The spicules are similar in size to the tetralophs. (c) Rare, thin triradiates and tetraradiates,<br />

10-17 JJL in total diameter (15 spicules). Some or all <strong>of</strong> the rays are terminally<br />

branched. The rays are only 1 p in basal diameter. The spicules may be developmental<br />

stages <strong>of</strong> the tetralophs and trilophs.<br />

DISCUSSION. The species is characterized by the lack <strong>of</strong> a special category <strong>of</strong> heterolophose<br />

candelabra. In addition, modified calthrops are absent. Two other species <strong>of</strong><br />

Corticium occur in the West Indies. C. versatile Schmidt, 1880, has peculiar candelabras<br />

and variously modified calthrops. C. quadripartitum Topsent, 1923, has special dermal<br />

heterolophose candelabras. Its tetralophs are similar in form to those <strong>of</strong> the Jamaican<br />

sponge but are slightly larger in size.<br />

DISTRIBUTION BY HABITAT<br />

The sponge fauna <strong>of</strong> individual habitats can now be summarized.<br />

(A) Port Royal Docks: The common species are:<br />

Ircinia fasciculata Halichondria melanadocia<br />

Ircinia strobilina Tedania ignis<br />

Darwinella rosacea Microciona microchela<br />

Desmapsamma anchorata Microciona rarispinosa<br />

Iotrochota birotulata Mycale laevis<br />

Gelliodes areolata<br />

The following species were present but less common:<br />

Verongia fistularis Placospongia carinata<br />

Dysidea fragilis Zygomycale parishii


78 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

Sponges were the dominant members <strong>of</strong> the sessile seawall community, which<br />

also included mussels, barnacles and ascidians. The species exhibited no obvious<br />

zonation on the piers and seawalls. Very large specimens <strong>of</strong> Ircinia fasciculata,<br />

Ircinia strobilina and Verongia fistularis were restricted to the base <strong>of</strong> the seawall.<br />

At that depth (8-10 ft.), sponges are certainly less subject to the surface salinity<br />

changes noted by Goodbody (1961) after heavy rains.<br />

(B) Port Royal Harbor:<br />

Verongia longissima<br />

Spheciospongia vesparia<br />

(C) Rasta's Wreck:<br />

Iotrochota birotulata Tedania ignis<br />

Sigmadocia caerulea Microciona microchela<br />

Halichondria melanadocia Microciona rarispinosa<br />

Halichondria magniconulosa Mycale laevis<br />

Sponges and ascidians were the most conspicuous elements <strong>of</strong> the wreck fauna.<br />

The microcionids incrusted mussel shells growing on the wreck. Microciona<br />

microchela covered large areas <strong>of</strong> the timbers as a thin film. Halichondria magniconulosa<br />

was found only in this habitat.<br />

(D) Port Royal turtle grass bed:<br />

Haliclona doria Mycale microsigmatosa<br />

Haliclona erina Tethya seychellenis<br />

Mycale microsigmatosa and Tethya seychellensis grew on turtle grass blades.<br />

The other two species <strong>of</strong> Tethya at Port Royal were restricted to the rocky<br />

environment <strong>of</strong> the cays. Haliclona doria formed repent colonies over the mud<br />

and turtle grass with most <strong>of</strong> the branches above the level <strong>of</strong> the substratum.<br />

Most specimens <strong>of</strong> Haliclona erina grew at the border between turtle grass and<br />

sand, near Rasta's wreck.<br />

(E) Mangroves bordering inlet:<br />

Dysidea fragilis Tedania ignis<br />

Sigmodocia caerulea Mycale laevis<br />

Halichondria melanadocia<br />

Sponges and ascidians covered many mangrove roots.<br />

(F) Main Boat Channel: The common species were:<br />

Darwinella rosacea Ulosa hispida<br />

Haliclona hogarthi Terpios zeteki<br />

Adocia implexiformis Geodia (Cydonium) papyracea<br />

Pellina coeliformis Geodia (Geodia) gibberosa<br />

Halichondria melanadocia<br />

Less common species were:<br />

Ircinia fasciculata Haliclona erina<br />

Dysidea fragilis Mycale microsigmatosa<br />

Sponges and ascidians covered a majority <strong>of</strong> the mangrove roots bordering the<br />

channel. Most species grew only on the root systems. A few specimens <strong>of</strong> Geodia


DISTRIBUTION BY HABITAT 79<br />

gibberosa and Terpios zeteki were found in the stream bed. Pellina coeliformis<br />

and a few specimens <strong>of</strong> Haliclona erina grew over the surface <strong>of</strong> Geodia gibberosa.<br />

A single root system <strong>of</strong>ten served as the substratum for several species <strong>of</strong><br />

sponge. The species exhibited no obvious horizontal or vertical zonation.<br />

The channel sponges were unaffected by the moderate autumn rains <strong>of</strong> 1961.<br />

Heavy seasonal rains are known to produce a surface layer <strong>of</strong> low salinity water<br />

in Kingston Harbor which is fatal to sponges (cf. Goodbody, 1961).<br />

The mangrove channel fauna has a sharply limited distribution at Port Royal.<br />

Six <strong>of</strong> the nine common species were growing exclusively in the mangroves, and<br />

none were found at the cays. Three <strong>of</strong> the five common species <strong>of</strong> habitat E did<br />

not extend into the channels.<br />

(G) Pigeon House Pile:<br />

Ircinia fasciculata<br />

Oligoceras hemorrhages<br />

Darwinella rosacea<br />

Haliclona erina<br />

(H) Shallow water at the cays:<br />

Ircinia fasciculata<br />

Ircinia strobilina<br />

Ne<strong>of</strong>ibularia massa<br />

Adocia albifragilis<br />

Acanthacarnus souriei<br />

Spirastrella coccinea<br />

Anthosigmella varians<br />

Diplastrella megastellata<br />

Cliona vermifera<br />

Most species, including Ne<strong>of</strong>ibularia massa, were incrustations growing on the<br />

underside <strong>of</strong> rocks. The heavy surf may exclude most massive species. Twelve<br />

<strong>of</strong> the seventeen species were collected only in this habitat. Only three species<br />

were also found in shallow water at Port Royal.<br />

(I) Deep water near cays:<br />

Ircinia fasciculata<br />

Ircinia strobilina<br />

Verongia fistularis<br />

(Ianthella ardis, in<br />

St. Thomas Parish.)<br />

Haliclona rub ens<br />

Desmapsamma anchorata<br />

Anthosigmella varians<br />

Placospongia carinata<br />

Geodia (Geodia) gibberosa<br />

Cliona viridis<br />

Placospongia carinata<br />

Tethya actinia<br />

Tethya sp.<br />

Erylus ministrongylus<br />

Chondrilla nucula<br />

Plakortis zyggompha<br />

Corticium tetralophum<br />

Ne<strong>of</strong>ibularia massa<br />

Iotrochota birotulata<br />

Gailyspongia fallax<br />

Callyspongia vaginalis<br />

Callyspongia plicifera<br />

Thalyseurypon conulosa<br />

Six <strong>of</strong> the species were restricted to this habitat. Most specimens <strong>of</strong> Ne<strong>of</strong>ibularia<br />

massa were distinctly vasiform, with upright growth, rather than thickly incrusting.<br />

(J) Offshore turtle grass beds:<br />

Oligoceras hemorrhages<br />

Dysidea fragilis<br />

Adocia carbonaria<br />

Sigmadocia caerulea<br />

Tedania ignis<br />

Lissodendoryx isodictyalis<br />

Tethya seychellensis


80 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

The <strong>of</strong>fshore turtle grass beds have a sponge fauna quite unlike that <strong>of</strong> the Port<br />

Royal turtle grass area. The <strong>of</strong>fshore beds are sandy rather than muddy and subject<br />

to considerable surf. The sponges grew at the base <strong>of</strong> the turtle grass clumps<br />

and on stones; none grew on the turtle grass blades. Adocia carbonaria and Lissodendoryx<br />

isodictyalis were found only in this habitat.<br />

FIELD KEY TO THE SPONGES OF PORT ROYAL<br />

The key is designed for use as a preliminary guide in field work. It is based<br />

on the appearance in life <strong>of</strong> typical members <strong>of</strong> each species. The key must be<br />

used with caution, and all identifications should be confirmed by comparison<br />

with the description. Undescribed species may be found, particularly at the<br />

outer cays and in slightly deeper water. Interesting specimens should be preserved<br />

in alcohol, after completing field notes, and sent to the author.<br />

The color in life is used for an initial division.<br />

A—Black and grayish-black.<br />

B—Blue and blue-green.<br />

C—Brown or tan.<br />

D—Gray.<br />

E—Green.<br />

F—Orange.<br />

G—Pink.<br />

H—Purple.<br />

I—Red and red-orange.<br />

J—White and <strong>of</strong>f-white.<br />

K—Yellow.<br />

Glossary <strong>of</strong> terms:<br />

Conule—a spike-shaped, thorn-like projection.<br />

Oscule—a large exhalant opening on the surface.<br />

(A) Near black and grayish-black.<br />

1—Conulose—2.<br />

1—Not conulose—3.<br />

2—Conules 3-7 mm high; massive; fetid—Ircinia strobilina.<br />

2—Conules less than 1 mm high; vasiform—Verongia fistularis (Turns black<br />

soon after collection; in life, has a yellow color.)<br />

3—Old specimens much larger than fist-sized; with clusters <strong>of</strong> oscules—Spheciospongia<br />

vesparia.<br />

3—Fist-sized or smaller; without clusters <strong>of</strong> oscules—4.<br />

4—Black; brittle; repent to lobate; stains alcohol blue-black—Adocia carbonaria.<br />

4—Near-black, with greenish tinge in interior; compressible; massive—Halichondria<br />

melanadocia.<br />

(B) Blue or blue-green.<br />

1—Conulose—Dysidea fragilis.<br />

1—Not conulose—2.<br />

2—Incrusting; blue-green, speckled with red-orange embryos—Mycale microsigmatosa.<br />

2—Massive or lobate—3.


FIELD KEY 81<br />

3—Tough; with skin-like surface; oscules flush with surface—Terpios zeteki.<br />

3—Compressible; surface not skin-like; with oscular projections or lobes—Sigmadocia<br />

caerulea.<br />

(C) Brown or tan.<br />

1—Conulose—2.<br />

1—Not conulose—3.<br />

2—Massive, usually larger than fist-sized; conules 0.5-3 mm high; fetid—Ircinia<br />

fasciculata.<br />

2—Small, lobate; conules less than 1 mm high; red exudate omitted upon handling<br />

after death—Oligoceras hemorrhages.<br />

3—Thinly incrusting—4.<br />

3—Not thinly incrusting—7.<br />

4—Surface with a stony armor <strong>of</strong> polygonal plates—Placospongia carinata.<br />

4—Surface unarmored—5.<br />

5—Consistency tough or rubbery—6.<br />

5—Consistency not tough or rubbery—Spirastrella coccinea or Diplastrella megastellata.<br />

6—Mottled glossy surface, with pale areas around oscules—Chondrilla nucula.<br />

6—Uniformly brown, not glossy—Plakortis zyggompha.<br />

7—Brittle or stiff—8.<br />

7—Not brittle or stiff-—9.<br />

8—Thickly incrusting to massive; stiff; dull brown—Anthosigmella varians.<br />

8—Report, cylindrical branches; brittle; dark brown—Haliclona doria.<br />

9—Easily crumbled; rough surface; massive or vasiform; irritating to the hands—<br />

Ne<strong>of</strong>ibularia massa.<br />

9—Tough; with skin-like surface—10.<br />

10—Dark brown; cortex removable in flakes—Erylus ministrongylus.<br />

10—Tan or brown; cortex not removable in flakes—Zygomycale parishii.<br />

(D) Gray.<br />

1—Conulose—2.<br />

1—Not conulose—Erylus ministrongylus.<br />

2—Tough; conules 3-7 mm high; fetid—Ircinia strobilina.<br />

2—S<strong>of</strong>t; conules 0.5-1 mm high—Dysidea fragilis.<br />

(E) Green.<br />

1—Spherical or hemispherical; tuberculate—some specimens <strong>of</strong> Tethya are colored<br />

green by algae; see under orange.<br />

1—Not spherical—2.<br />

2—Easily crumbled; without a skin; massive to ramose—Haliclona erina.<br />

2—Tough; surface skin-like; lobate to digitate—Terpios zeteki.<br />

(F) Orange.<br />

1—Spherical or hemispherical; tuberculate—2.<br />

1—Not spherical—3.<br />

2—Spherical; usually larger than 1 cm in diameter; on turtle grass—Tethya seychellensis.<br />

3—Incrusting—4.<br />

3—Not incrusting—5.<br />

4—Smooth speckled, with red-orange embryos—Mycale microsigmatosa.


82 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

4—Projecting spicules and fleshy tufts conspicuous; uniformly orange—Ulosa<br />

hispida.<br />

5—Compressible; with rimmed oscules—Mycale laevis.<br />

5—Tough; oscules flush with surface—Terpios zeteki.<br />

(G) Pink,<br />

1—Ramose—Desmapsamma anchorata.<br />

1—Incrusting—2.<br />

2—Conulose—Darwinella rosacea.<br />

2—Smooth—3.<br />

3—Tough; with pinkish-orange interior—Microciona microchela.<br />

3—S<strong>of</strong>t, speckled, with red-orange embryos—Mycale microsigmatosa.<br />

(H) Purple.<br />

1—Conulose—2.<br />

1—Not conulose—7.<br />

2—With only a few thorn-like conules on an otherwise smooth surface; dull<br />

purple—Callyspongia fallax.<br />

2—With numerous conules—3.<br />

3—Cheese-like consistency; bluish-purple—Ianthella ardis.<br />

3—Characters otherwise—4.<br />

4—Very dark purple—5.<br />

4—Light purple—6.<br />

5—Dark reddish purple—Thalyseurypon conulosa.<br />

5—Nearly black, with a greenish tinge in life; orange zoanthids on surface—<br />

Iotrochota birotulata.<br />

6—Tough; reddish-purple; fetid—Ircinia fasciculata.<br />

6—S<strong>of</strong>t; pale purple—Dysidea fragilis.<br />

7—Vasiform—8.<br />

7—Not vasiform—9.<br />

8—With a paper-thin terminal collar—Callyspongia plicifera.<br />

8—Without a collar—Callyspongia vaginalis.<br />

9—Tough; with a skin-like surface—10.<br />

9—Characters otherwise.<br />

10—Lobate or digitate; on mangroves—Terpios zeteki.<br />

10—Usually with elongate branches; on pilings—Zygomycale parishii.<br />

11—Massive, with a fetid odor—Lissodendoryx isodictyalis.<br />

11—Ramose or with oscular projections—12.<br />

12—Easily crumbled, with oscular projections—Adocia implexiformis.<br />

12—Not easily crumbled, cylindrical branches, oscules flush with surface—13.<br />

13—Limp; an anastomosing repent complex <strong>of</strong> branches—Haliclona hogarthi.<br />

13—Compressible; with a very rough surface; with stout, flattened erect branches<br />

—Gelliodes areolata.<br />

(I) Red-orange and red.<br />

1—Conulose—2.<br />

1—Not conulose—3.<br />

2—Massive; reddish-purple; fetid odor—Ircinia fasciculata.<br />

2—Cylindrical; dull red—Verongia longissima (Yellow in life, changing soon<br />

after collection.)<br />

3—Spherical; tuberculate—Tethya seychellensis.


ZOOGEOGRAPHICAL NOTES 83<br />

3—Not spherical—4.<br />

4—Boring, with papillae at surface <strong>of</strong> substratum—Cliona vermifera.<br />

4—Not boring—5.<br />

5—Incrusting—6.<br />

5—Not incrusting—7.<br />

6—Punctiform surface; on shells and pilings at Port Royal—Microciona rarispinosa.<br />

6—Surface not punctiform; on rocks at cays—Acanthacarnus souriei (Spirastrella<br />

coccinea has been reported as red in the literature.)<br />

7—Massive or lobate; red-orange; with conspicuous red-orange gemmules in the<br />

interior—Tedania ignis.<br />

7—Characters otherwise-—8.<br />

8—With a skin-like surface—Terpios zeteki.<br />

8—Without a skin-like surface—9.<br />

9—Dark red, retaining that color in preservative—Haliclona rubens.<br />

9—Pale color, dull red or reddish-purple—10.<br />

10—Reddish-purple; limp, a repent complex <strong>of</strong> branches—Haliclona hogarthi.<br />

10—Dull red; compressible; with stout flattened erect branches; with a rough surface—Gelliodes<br />

areolata.<br />

(J) White and <strong>of</strong>f-white.<br />

1—Massive—2.<br />

1—Incrusting—3.<br />

2—Leathery to stony; with a thick rigid cortex; with conspicuous sieve areas—<br />

Geodia gibber osa.<br />

2—Easily crushed; with a paper-like cortex; without oscular areas—Geodia<br />

papyracea.<br />

3—Oscules flush with the surface—Adocia albifragilis.<br />

3—Oscules on high paper-thin tubes—Pellina coeliformis.<br />

(K) Yellow.<br />

1—Conulose—2.<br />

1—Not conulose—4.<br />

2—Vasiform—Verongia fistularis.<br />

2—Not vasiform—3.<br />

3—Cylindrical—Verongia longissima.<br />

3—Massive—Ianthella ardis.<br />

4—Boring sponges with papillae at the surface <strong>of</strong> the substratum—Cliona viridis.<br />

4—Not boring—5.<br />

5—Incrusting—Corticium tetralophum.<br />

5—Not incrusting—6.<br />

6—Lobate, s<strong>of</strong>t—Callyspongia pallida.<br />

6—Massive, tough—Halichondria magniconulosa.<br />

ZOOGEOGRAPHICAL NOTES<br />

According to present records, most Jamaican sponges have a limited distribution.<br />

Of the 57 recorded species, (including three not found in the present study),<br />

18 have not been collected elsewhere. An additional <strong>20</strong> species, including the<br />

partially classified one, are known only from tropical Atlantic American waters.<br />

About two-thirds <strong>of</strong> the Jamaican fauna (38 species) are therefore endemic to the


84 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

region. Five other species, Ircinia fasciculata, 7. strobilina, Verongia lacunosa,<br />

Adocia carbonaria, and Tedania ignis, may or may not be conspecific with morphologically<br />

similar sponges in other parts <strong>of</strong> the world.<br />

Ten species are widely distributed outside the West Indies. Dysidea fragilis,<br />

Lissodendoryx isodictyalis, Cliona viridis, and Chondrilla nucula are circumtropical<br />

or nearly so, and the first three have also been reported from temperate<br />

waters. Other widespread species are Desmapsamma anchorata (West Africa,<br />

Indo-Pacific), Zygomycale parishii (Indo-Pacific), Spirastrella coccinea (Tropical<br />

Pacific America, West Africa, Mediterranean-Atlantic), Cliona vermifera (Mediterranean,<br />

Central Pacific), Placospongia carinata (Indo-Pacific, tropical Pacific<br />

America), and Tethya seychellensis (Indo-Pacific). The disjunct patterns indicate<br />

the fragmentary state <strong>of</strong> our present knowledge. In addition, several <strong>of</strong> the<br />

supposed species, such as Lissodendoryx isodictyalis and Cliona viridis, may prove<br />

to be complexes <strong>of</strong> closely related species.<br />

Of the remaining five species, Tethya actinia has been recorded from the West<br />

Central Pacific. Acanthacarnus souriei has been previously recorded only from<br />

West Africa and the Mediterranean. Geodia gibberosa and Mycale microsigmatosa<br />

also occur in West Africa. The former species has been collected at the<br />

Pacific end <strong>of</strong> the Panama canal. The latter species has been recorded from the<br />

Pacific coast <strong>of</strong> Panama and from the Hawaiian Islands. Terpios zeteki is known<br />

from both coasts <strong>of</strong> Panama, the Gulf <strong>of</strong> California and the Hawaiian Islands.<br />

The coasts <strong>of</strong> Central America require careful distributional studies. Several<br />

other West Indian species are found on the Pacific side <strong>of</strong> Panama (cf. de Laubenfels,<br />

1936b) and a few reach the Gulf <strong>of</strong> California, (cf. Dickinson, 1945).<br />

De Laubenfels (1950b, 1951a) recorded a number <strong>of</strong> species which are common<br />

to the West Indies and Hawaii. The Jamaican specimens <strong>of</strong> Mycale microsigmatosa<br />

and Terpios zeteki agree closely with his Hawaiian descriptions. The<br />

presence <strong>of</strong> a considerable number <strong>of</strong> species common to both sides <strong>of</strong> the Isthmus<br />

<strong>of</strong> Panama, a barrier in existence since the Pliocene, raises the possibility <strong>of</strong><br />

transport by ship in very recent times. De Laubenfels indeed found a Panamanian<br />

sponge, presumably Mycale cecilia = M. microsigmatosa, on dry-docked ships at<br />

Pearl Harbor (cf. de Laubenfels, 1950b, p. 4).<br />

The present study suggests that the Jamaican shallow-water fauna has close<br />

affinities with those <strong>of</strong> the southern Caribbean. Nine species have been previously<br />

recorded only from that little-known area. Burton (1930) has suggested that the<br />

distribution <strong>of</strong> sponges is closely related to ocean currents which can transport<br />

larvae. Jamaica lies in the path <strong>of</strong> surface currents which run from the Lesser<br />

Antilles and the Venezuelan coast toward the Yucatan Channel. Only a few Jamaican<br />

species have been recorded from Bermuda and North Carolina which<br />

are rather distant from Jamaica in terms <strong>of</strong> the paths <strong>of</strong> ocean currents. Burton's<br />

theory could be tested further by studying the sponge fauna <strong>of</strong> the Lesser<br />

Antilles; a strong West African element would be expected through the transport<br />

<strong>of</strong> larvae by the equatorial currents.<br />

SUMMARY<br />

A total <strong>of</strong> 54 species have been recorded in a systematic study <strong>of</strong> the shallow<br />

water sponges <strong>of</strong> Port Royal, Jamaica. The new genus Ne<strong>of</strong>ibularia is proposed<br />

for the preoccupied name Fibularia Carter. Sixteen <strong>of</strong> the species are new to<br />

science, namely: Darwinella rosacea, Haliclona hogarthi, Adocia implexiformis,<br />

Adocia albifragilis, Pellina coeliformis, Sigmadocia caerulea, Callyspongia pallida,


LITERATURE CITED 85<br />

Halichondria magniconulosa, Microciona microchela, Microciona rarispinosa,<br />

Thalyseurypon conulosa, Ulosa hispida, Diplastrella megastellata, Geodia (Cydonium)<br />

papyracea, Erylus ministrongylus, and Corticium tetralophum.<br />

LITERATURE CITED<br />

References through 1913 are dated in accordance with the bibliography <strong>of</strong> Vosmaer<br />

and can be found in that volume. The letters a, b, c, etc., have been employed in place<br />

<strong>of</strong> the alpha, beta, gamma system used by Vosmaer. Pre-1913 references, when listed by<br />

Vosmaer, are not included in the present bibliography.<br />

Pre-1913 references:<br />

Vosmaer, G. C. J., 1923. Bibliography <strong>of</strong> sponges, 1551-1913. Cambridge Univ. Press,<br />

London. 234 p.<br />

Later references, and references not listed by Vosmaer:<br />

Alander, Harald, 1942. Sponges from the Swedish West coast and adjacent waters. Goteborg,<br />

Henrik Struves. 1-95.<br />

Annandale, Nelson, 1915. Indian boring sponges <strong>of</strong> the family Clionidae. Rec. Indian<br />

Mus., vol. 11 (1): 1-24.<br />

Arndt, Walther, 1927. Kalk-und Kieselschwamme von Curasao. Bijdr. Dierk. Amsterdam,<br />

vol. 25: 133-158.<br />

1943. Die tiergeographische Gliederung der Schwammfauna der Nord- und Ostsee.<br />

Arch. Naturgesch., ser. 2, vol. 12: 303-349.<br />

Babic\ K., 1922. Monactinellida und Tetractinellida des Adriatischen Meeres. Zool. Jb.<br />

(Syst. Geogr., und Biol.), Vol. 46: 217-302.<br />

Bergquist, P. R., 1961. The Keratosa (Porifera) collected by the Chatham Islands 1954<br />

expedition. Bull. N.Z. Dep. sci. industr. res., vol. 139 (5): <strong>20</strong>7-219.<br />

Br0ndsted, H. V., 1934. R&ultats scientifiques du voyage aux Indes Orientales neerlandaises<br />

de LL. AA. RR. le Prince et la Princesse Leopold de Belgique. Sponges. Mem.<br />

Mus. Hist. nat. Belg., vol. 2 (15): 3-27.<br />

Burton, Maurice, 1924a. A revision <strong>of</strong> the sponge family Donatiidae. Proc. zool. Soc.<br />

London: 1033-1045.<br />

1924b. The genus Chondrilla. Ann. Mag. nat. Hist., Vol. (9) 14: <strong>20</strong>6-<strong>20</strong>9.<br />

1929. Porifera II. Antarctic Sponges. Nat. Hist. Rep. Terra Nova Exped. Zool.,<br />

vol. VI (4): 393-458. Br. Mus. (Nat. Hist.)<br />

1930. Norwegian sponges from the Norman collection. Proc. zool. Soc. London:<br />

487-546.<br />

1932. Sponges. 'Discovery* Rep., vol. 6: 237-392. Brit. Mus. (Nat. Hist.).<br />

1933. Report on a small collection <strong>of</strong> sponges from Stil Bay, South Africa. Ann.<br />

Mag. nat. Hist., vol. 11: 235-244.<br />

1934. Sponges. Sci. Rep. Gr. Barrier Reef Exped., vol. IV (14): 513-621. Brit. Mus.<br />

(Nat. Hist.)<br />

1936a Sponges. The fishery grounds near Alexandria. Notes Fish. Res. Div. Cairo,<br />

vol. 17: 1-28.<br />

1936b. Notes on sponges from South Africa, with descriptions <strong>of</strong> new species.<br />

Ann. Mag. nat. Hist., vol. 17: 141-147.<br />

1937. The littoral fauna <strong>of</strong> Krusadai Island in the Gulf <strong>of</strong> Manaar. Bull. Madras<br />

Govt. Mus., N.S., Nat. Hist., vol. 1-2 (4): 1-58.<br />

1948. Marine sponges <strong>of</strong> the Congo coast. Bull. Inst. roy. Colon. Beige, vol. 19 (3):<br />

750-752.<br />

1954. The "Rosaura" expedition, 5. Sponges. Bull. Brit. Mus. (Nat. Hist.) Zool.,<br />

vol. 2 (6): 215-239.<br />

1956. The sponges <strong>of</strong> West Africa. Atlantide Rpt. vol. 4: 111-147. Danish Science<br />

Press, Copenhagen.<br />

1959a. Sponges. Sci. Rep. Murray Exped., vol. 10 (5): 151-281. Brit. Mus. (Nat.<br />

Hist.)


86 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

1959b. Sponges. Zool. Iceland, vol. 2 (3-4): 1-71.<br />

Burton, Maurice and Rao, H. S., 1932. Report on the shallow water marine sponges in<br />

the collection <strong>of</strong> the Indian <strong>Museum</strong>. Rec. Indian Mus. vol. 34: 299-358.<br />

De Laubenfels, M. W., 1932a. Physiology and morphology <strong>of</strong> Porifera exemplified by<br />

Iotrochota birotulata Higgin. Carnegie Institution <strong>of</strong> Washington Publication No.<br />

435: 37-66.<br />

1932b. The marine and fresh water sponges <strong>of</strong> California. Proc. U. S. nat. Mus.,<br />

vol. 81 (4): 1-140.<br />

1934. New sponges from the Puerto Rican deep. Smithson. Misc. Coll., vol. 91,<br />

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1935a. Some sponges <strong>of</strong> Lower California (Mexico). Amer. Mus. Novit. 779: 1-14.<br />

1935b. A collection <strong>of</strong> sponges from Puerto Galera, Mindora, Philippine Islands.<br />

Philipp. J. Sci., vol. 56 (3): 327-336.<br />

1936a. Sponge fauna <strong>of</strong> the Dry Tortugas. Carnegie Inst, <strong>of</strong> Washington Publication<br />

No. 467: 1-225.<br />

1936b. A comparison <strong>of</strong> the shallow water sponges near the Pacific end <strong>of</strong> the<br />

Panama canal with those at the Caribbean end. Proc. U. S. nat. Mus., vol. 83, (2993):<br />

441-466.<br />

1939. Sponges collected on the Presidential Cruise <strong>of</strong> 1938. Smithson. Misc. Coll.,<br />

vol. 98 (15): 1-7.<br />

1947. Ecology <strong>of</strong> the sponges <strong>of</strong> a brackish water environment at Beaufort, N.C.<br />

Ecol. Monogr., vol. 17: 31-46.<br />

1948. The order <strong>of</strong> Keratosa <strong>of</strong> the Phylum Porifera. A monographic study. Occ.<br />

Pap. Allan Hancock Fdn. No. 3: 1-217.<br />

1949. Sponges <strong>of</strong> the Western Bahamas. Amer. Mus. Novit. no. 1431: 1-25.<br />

1950a. The Porifera <strong>of</strong> the Bermuda Archipelago. Trans, zool. Soc. Lond., vol. 27<br />

(1): 1-<strong>20</strong>1.<br />

1950b. Sponges <strong>of</strong> Kaneohe Bay. Pacif. Sci., vol. IV (1): 3-36.<br />

. 1951a. Sponges <strong>of</strong> the island <strong>of</strong> Hawaii. Pacif. Sci., vol. V. (3): 256-271.<br />

1951b. A collection <strong>of</strong> sponges from the Black Sea. Arch. Hydrobiol. Bd. XLV:<br />

213-216.<br />

1953a. Sponges <strong>of</strong> the Gulf <strong>of</strong> Mexico. Bull. Mar. Sci. Gulf Caribbean, vol. 2 (3):<br />

511-557.<br />

1953b. A guide to the sponges <strong>of</strong> Eastern North America. Special Publ. U. <strong>of</strong><br />

Miami: 1-32.<br />

« 1954a. The sponges <strong>of</strong> the West-central Pacific. Oregon State Monogr. Zool., no. 7:<br />

1-306.<br />

1954b. Occurrence <strong>of</strong> sponges in an aquarium. Pacif. Sci., vol. VIII (3): 337-340.<br />

1955. Sponges <strong>of</strong> Onotoa. Pacif. Sci., vol. IX (2): 137-143.<br />

1956. Preliminary discussion <strong>of</strong> the sponges <strong>of</strong> Brazil. Oceanographica Biologica<br />

Universidade de Sao Paulo, No. 1: 1-4.<br />

1957. New species and records <strong>of</strong> Hawaiian sponges. Pacif. Sci., vol. XI (2) 236-<br />

251.<br />

Dendy, Arthur, 1916a. Report on the non-calcareous sponges collected by Mr. James<br />

Hornell at Okhamandal in Kattiawar in 1905-6. In: Hornell, J., Report to the government<br />

<strong>of</strong> Baroda on the Marine Zoology <strong>of</strong> Okhamandal in Kattiawar. Vol. 2: 93-146.<br />

1916b. Report on the Homosclerophora and Astrotetraxonida collected by H. M. S.<br />

Sealark in the Indian Ocean. Trans. Linn. Soc. Lond. (Zool.), vol. 17: 225-271.<br />

1921. Report on the Sigmatotetraxonida collected by H. M. S. Sealark in the<br />

Indian Ocean. Trans. Linn. Soc. Lond. (Zool.), vol. 18: 1-164.<br />

1924. Porifera 1. Non-Antarctic Sponges. Nat. Hist. Rep. Terra Nova Exped.,<br />

Zool., vol. 6 (3): 269-392. Brit. Mus. (Nat. Hist.)<br />

Dendy, Arthur and Frederick, L. M., 1924. On a collection <strong>of</strong> sponges from Abrolhos<br />

Islands, West Australia. J. Linn. Soc. London, vol. 35: 477-518.<br />

Dickinson, M. G., 1945. Sponges <strong>of</strong> the Gulf <strong>of</strong> California. Allan Hancock Pacif. Exped.,<br />

vol. 11 (1): 1-251.


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Goodbody, I. M., 1961. Mass mortality <strong>of</strong> a marine fauna following tropical rains. Ecology,<br />

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George, W. C. and Wilson, H. V., 1919. Sponges <strong>of</strong> Beaufort (N.C.) harbor and vicinity.<br />

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Hartman, W. D., 1955. A collection <strong>of</strong> sponges from the West Coast <strong>of</strong> the Yucatan<br />

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1958. <strong>Natural</strong> <strong>History</strong> <strong>of</strong> the Marine Sponges <strong>of</strong> Southern New England. Bull.<br />

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Hentschel, Ernst, 1923. Porifera. In: Handbuch der Zoologie, von W. Kukenthal. Berlin.<br />

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1929. Die Kiesel-und Hornschwamme des Nordlichen Eismeers. Fauna Arct., vol.<br />

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1917. Esponjas del litoral de Asturias. Trab. Mus. cienc. nat., Zool., vol. 36: 5-39,<br />

Hopkins, S. H., 1956. The boring sponges which attack South Carolina Oysters, with<br />

notes on some associated organisms. Contr. Bears Bluff Lab., vol. 23: 3-30.<br />

Kumar, A., 1925. Report on some tetraxonid sponges in the collection <strong>of</strong> the Indian<br />

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Levi, Claude, 1952. Spongiaires de la cote du S£n£gal. Bull. Inst, franc,. Afr. Noire, vol.<br />

14 (1): 34-59.<br />

1953. Description de Plakortis nigra n. sp. et remarques sur les Plakinidae. (Demosponges).<br />

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1956a. Spongiaires des cotes de Madagascar. M&n. Inst. sci. Madagascar, vol. (4)<br />

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1956b. £tude des Halisarca de Rosc<strong>of</strong>f. Embryologie et Systematique des De*mosponges.<br />

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1957. Spongiaires des cotes d'Israel. Bull. Res. Council Israel, vol. 6 (B) (3-4): <strong>20</strong>1-<br />

212.<br />

1959. Spongiaires. Result, sci. Camp. "Calypso," fasc. 4, Ann. Inst, oceanogr.<br />

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McDougall, K. D., 1943. Sessile marine invertebrates <strong>of</strong> Beaufort, N.C. Ecol. Monogr.,<br />

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88 A SYSTEMATIC STUDY OF THE DEMOSPONGIAE OF JAMAICA<br />

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100 (2): 273-532.


SYSTEMATIC INDEX<br />

Numbers in boldface refer to detailed descriptions <strong>of</strong> species; numbers in italics refer to text<br />

illustrations; plate references are in capital Roman numerals.<br />

Acanthacarnus, 40; levii, 40; souriei, 7, 40, 79,<br />

83-84<br />

acanthifolium, Pandaros, 45<br />

acerata, Iotrochota, 24<br />

aceratoobtusa, Microciona, 44<br />

actinia, Tethya, 7, 66-67, 79, 84<br />

Adocia, 26, 27; albifragilis, vi, 1, 6, 28-29, 28,<br />

79, 83-84; baeri, 28; carbonaria, vi, 6, 26-27,<br />

79-80, 84, III; chilensis, 28; grossa, 28; implexiformis,<br />

vi, 1, 6, 27-28, 27, 78, 82, 84, II;<br />

neens, 29; tubifera, 28<br />

Adociidae, v, 1, 5, 6, 26<br />

albifragilis, Adocia, vi, 1, 6, 28-29, 28, 79, 83-84<br />

Alcyonium, purpureum, 54; vesparium, 57<br />

Alemo seychellensis, 65-66<br />

alleni, Erylus, 74<br />

Amphilectidae, 1, 5, 45<br />

Amphimedon arborescens, 19<br />

anchorata, Desmapsamma, vi, 6, 21-22, 77, 79,<br />

82, 84, II; Fibularia, 21-22<br />

andrewsii, Spirastrella, 4, 57, 58<br />

angulata, Hymeniacidon, 61-62<br />

angulosa, Spongia, 51<br />

angulospiculata, Epipolasis, 4<br />

angulospiculatus, Suberites, 4, 6<br />

anhelans, Tedania, 37-38<br />

Anthosigmella, 55, 57; coronarius, 4, 56; varians,<br />

vi, 4, 7, 55-57, 79, 81, VII<br />

Aplysillidae, v, 6, 17<br />

Aplysina, 14; cauliformis, 13-14; fenestrata, 13;<br />

fistularis, 12; flagelliformis, 4, 13-14; flagelliformis<br />

rugosa, 13; hirsuta, 12; longissima, 13;<br />

praetexta, 13; spengelii, 4<br />

arborescens, Amphimedon, 19<br />

arcuata, Papillina, 56-57<br />

ardw, Ianthella, 6, 16-17, 79, 82-83<br />

areolata, Gelliodes, vi, 5-6, 25-26, 77, 82-83, II;<br />

Halichondria, 15; Haliclona, 25; Pachychalina,<br />

25<br />

aspinosa, Callyspongia, 37<br />

atlantica, Poterion, 57-58<br />

aurantiaca, Pachychalina, 26<br />

aurea, Verongia, 14<br />

australiensis, Chondrilla, 74-75<br />

Axocielita, 44; kilauea, 44; linda, 44<br />

Axociella, 43-44; cfl/Ja, 44<br />

6


conulosa, Thalyseurypon, vi, 1, 7, 44-45, 44, 79,<br />

82, 85, VI<br />

copiosa, Clathria, curacaoensis, 42<br />

coriacea, Dysidea, 15<br />

coronaria, Cliona, 56; Spirastrella, 56<br />

coronarius, Anthosigmella, 4, 56<br />

Corticiidae, 76<br />

Corticium, 77; quadripartitum, 77; tetralophum,<br />

vi, 1, 8, 77, 77, 79, 83, 85; versatile, 77<br />

crassissima, Mycale, 51<br />

Cribrella labiata, 61<br />

cribrosa, Papillina, 57<br />

cunctatrix, Spirastrella, 54-55; Spirastrella, porcata,<br />

55; Spirastrella, robusta, 55<br />

Cydonium, 70-72<br />

Darwinella, 17-18; intermedia, 18; joyeuxi, 18;<br />

miilleri, 18; rosacea, 1, 6, 17-18, 77-79, 82, 84<br />

decumbens, Spirastrella, 54-55<br />

dendroides, Ircinia, 10<br />

Desmacidon, carterianum, 21-22; reptans, 21-22<br />

Desmacidonidae, v, 6, 21, 45<br />

Desmapsamma, 21-22; anchorata, vi, 6, 21-22,<br />

77, 79, 82, 84, II<br />

digitata, Reniera, 37-38; Tedania, 37-38; Tecfama,<br />

bermudensis, 37-38<br />

Diplastrella, 58-59; bistellata, 54; megastellata,<br />

vi, 1, 7, 58-59, 5S, 79, 81, 85, VII; ornata, 59<br />

diploderma, Tethya, 66-67<br />

Donatia, ingalli seychellensis, 65; seychellensis,<br />

65-66; viridis, 65-66<br />

doria, Haliclona, vi, 6, 18, 78, 81, I<br />

Duseideia fragilis, 14-16<br />

Dysidea, 14; coriacea, 15; etheria, 16; fragilis, 6,<br />

14-16, 77-82, 84<br />

Dysideidae, v, 6, 14<br />

dysoni, Geodia, 70; Spongia, 57-58<br />

ect<strong>of</strong>ibrosa, Hircinia, 8-9<br />

Ectyoninae, 45<br />

elastica, Spongelia, lobata, 15-16; Spongelia,<br />

lobosa, 15-16; Spongelia, massa, 15-16<br />

embolophora, Chondrilla, 74-75<br />

Epipolasis angulospiculata, 4<br />

erina, Haliclona, vi, 6, 19-<strong>20</strong>, 78-79, 81, I<br />

Erylus, 72; alleni, 74; euastrum, 74; ministrongylus,<br />

vi, 1, 7, 72-74, 75, 79, 81, 85, VIII;<br />

transiens, 74<br />

eschrichtii, Callyspongia, 32<br />

Esperella, 45; parishii, 48, 50; plumosa, 48, 50;<br />

ridleyi, 48, 50<br />

Esperellinae, 45<br />

Esperia, 45; laevis, 46-47; parishii, 48, 50;<br />

plumosa, 48, 50<br />

Esperiadae, 45<br />

etheria, Dysidea, 16<br />

euastrum, Erylus, 74<br />

excelsa, Haliclona, 25<br />

/a/ZflX, Callyspongia, vi, 6, 31-32, 35, 79, 82, IV<br />

fasciculata, Hircinia, 8-9; Ircinia, 4, 6, 8-10,<br />

77-79, 81-82, 84; Spongia, 8-9<br />

INDEX 91<br />

favosa, Clathria, 42<br />

fenestrata, Aplysina, 13<br />

Fibularia, 1, 22-23, 84; anchorata, 21-22; mawfl,<br />

22-23; ramosa, 22<br />

fibulata, Gelliodes, 31<br />

fibulatus, Gellius, 31<br />

Fibulia, 22-23; bermuda, 23; carnosa, 22-23;<br />

massa, 23; nolitangere, 23; raphidifera, 23<br />

Fistularia fistularis, 12<br />

fistularis, Aplysina, 12; Fistularia, 12; Luffaria,<br />

12; Spongia, 12; Verongia, vi, 6, 12-13, 31,<br />

77-80, 83, I<br />

fistulata, Mycale, 48; Mycale, microsigmatosa,<br />

47-48<br />

flagelliformis, Aplysina, 4, 13-14; Aplysina,<br />

rugosa, 13; Hircinia, 14<br />

foliacea, Thalyseurypon, 45<br />

fragilis, Duseideia, 14-16; Dysidea, 6, 14-16,<br />

77-82, 84; Spongelia, 14-16; Spongia, 14-15<br />

Gelliodes, 22, 25; areolata, vi, 5-6, 25-26, 77,<br />

82-83, II; fibulata, 31; ramosa, 22<br />

Gellius, 31; fibulatus, 31; massa, 23<br />

Geodia, 19, 68, 70; cariboa, 69-70; carinata, 62,<br />

64; d;y5om, 70; gibberosa, 4, 7, 29, 68-70,<br />

78-79, 83-84, III, VlII; glariosus, 72; medsa,<br />

4, 69-70; papyracea, vi, 1, 7, 71-72, 71, 78, 83,<br />

85, VIII; reticulata, 69-70; tuberculosa, 69-70;<br />

tumulosa, 69-70<br />

Geodiidae, v, 7, 68, 71<br />

gibberosa, Geodia, 4, 7, 29, 68-70, 78-79, 83-84,<br />

III, VIII; Pyxto, 69-70<br />

gigantea, Hircinia, 10-11<br />

glariosus, Geodia, 72<br />

grossa, Adocia, 28<br />

Halichondria, 52-53; areolata, 15; birotulata, 4,<br />

24; bowerbanki, 53; isodictyalis, 38-39; magniconulosa,<br />

vi, 1, 7, 53, 55, 78, 83, 85, V;<br />

melanadocia, vi, 7, 52, 77-78, 80, V; panicea,<br />

53<br />

Halichondriidae, v, 7, 52<br />

Haliclona, 18; areolata, 25; doria, vi, 6, 18, 78,<br />

81, I; erma, vi, 6, 19-<strong>20</strong>, 78-79, 81, I; excelsa,<br />

25; hogarthi, vi, 1, 6, <strong>20</strong>-21, <strong>20</strong>, 78, 82-84, II;<br />

implexa, 28; longleyi, 18; rubens, 5-6, 18-19,<br />

79, 83; vmdw, <strong>20</strong><br />

Haliclonidae, v, 6, 18<br />

hemorrhages, Oligoceras, vi, 6, 11, 79, 81, I<br />

Hircinia, ect<strong>of</strong>ibrosa, 8-9; fasciculata, 8-9;<br />

flagelliformis, 14; gigantea, 10-11; strobilina,<br />

10-11; variabilis, 4, 8-9<br />

hirsuta, Aplysina, 12; Verongia, 13; Verongia,<br />

fistularoides, 12<br />

hispida, Ulosa, vi, 1, 7, 51-52, 51, 78, 82, 85<br />

hogarthi, Haliclona, vi, 1, 6, <strong>20</strong>-21, <strong>20</strong>, 78, 82-84,<br />

II<br />

Hymeniacidon, angulata, 61-62; pulvinatus,<br />

57-58<br />

lanthella, 16; ardis, 6, 16-17, 79, 82-83; fcasfa,<br />

16-17; ianthella, 17


92<br />

ianthella, Ianthella, 17<br />

ignis, Tedania, vi, 5, 7, 37-38, 39, 77-79, 83-84,<br />

V; Tedania, pacifica, 37; Thalysias, 37-38<br />

implexa, Haliclona, 28<br />

implexiformis, Adocia, vi, 1, 6, 27-28, 27, 78,<br />

82, 84, II<br />

incrustans, Spongelia, 15-16<br />

ingalli, Donatia, seychellensis, 65; Tethya, 65-67<br />

intermedia, Darwinella, 18; Placospongia, 62,<br />

64; Strongylacidon, 51<br />

Iotrochota, 24; acerata, 24; birotulata, 4-6,<br />

24-25, 77-79, 82; coccinea, 25<br />

Ircinia, 8-10, 39; dendroides, 10; fasciculata,<br />

4, 6, 8-10, 77-79, 81-82, 84; muscarum, 11;<br />

oro$, 10; ramosa, 9; strobilina, 6, 9, 10-11,<br />

77-81, 84; strobilina irregularis, 10-11; van"abilis,<br />

8-10<br />

isochela, Mycale, 48, 50, 51<br />

isodictyalis, Halichondria, 38-39; Lissodendoryx,<br />

7, 38-40, 79-80, 82, 84; Lissodendoryx, jacksoniana,<br />

38, 40; Lissodendoryx, paucispinosa,<br />

38, 40; Myxi7/a, 38-39<br />

jacksoniana, Lissodendoryx, 40<br />

joyeuxi, Darwinella, 18<br />

hilauea, Axocielita, 44<br />

labiata, Cribrella, 61<br />

lacunosa, Verongia, 4, 6, 13, 84<br />

laevis, Esperia, 46-47; Mycale, vi, 7, 46-47, 49,<br />

77-78, 82, VI<br />

Laxosuberites zeteki, 59-60<br />

leptoderma, Lissodendoryx, 38; Tedania, 38-39<br />

/ex/ft, Acanthacarnus, 40<br />

linda, Axocielita, 44<br />

linteiformis, Spongia, 10<br />

Lissodendoryx, 38; carolinensis, 39-40; isodictyalis,<br />

7, 38-40, 79-80, 82, 84; isodictyalis<br />

jacksoniana, 38, 40; isodictyalis paucispinosa,<br />

38, 40; jacksoniana, 40; leptoderma, 38;<br />

similis, 38-40<br />

longissima, Aplysina, 13; Verongia, 4, 6, 13-14,<br />

78, 82-83<br />

longleyi, Haliclona, 18<br />

Luffaria, 14; cauliformis, 14; fistularis, 12; sebae,<br />

13<br />

lyncurium, Tethya, 65<br />

magniconulosa, Halichondria, vi, 1, 7, 53, 55,<br />

78, 83, 85, V<br />

mflwfl, Fibularia, 22-23; Fibulia, 23; Gellius, 23;<br />

Ne<strong>of</strong>ibularia, 6, 23, 79, 81<br />

maunakea, Mycale, 48<br />

maunaloa, Microciona, 42<br />

maxima, Spinosella, 4, 34-35<br />

maza, Tethya, 7, 65-66, 67-68<br />

media, Geodia, 4, 69-70; Synops (?), 69<br />

megastellata, Diplastrella, vi, 1, 7, 58-59, 55, 79,<br />

81, 85, VII<br />

melanadocia, Halichondria, vi, 7, 52, 77-78,<br />

80, V<br />

INDEX<br />

melobesioides, Placospongia, 63-64<br />

membranacea, Ophlitaspongia, 44<br />

microchela, Microciona, vi, 1, 5, 7, 41-42, 42,<br />

77-78, 82, 85<br />

Microciona, 41-44; aceratoobtusa, 44; maunaloa,<br />

42; microchela, vi, 1, 5, 7, 41-42, 42, 77-78,<br />

82, 85; parthena, 42; rarispinosa, vi, 1, 7, 42-<br />

44, 45, 77-78, 83, 85; «m*7w, 43-44; spinosa, 43<br />

Microcionidae, v, 7, 41<br />

microsigmatosa, Mycale, 7, 47-48, 78, 80-82, 84<br />

millepora, Pachychalina, 26<br />

ministrongylus, Erylus, vi, 1, 7, 72-74, 75, 79,<br />

81, 85, VIII<br />

mixta, Placospongia, 62, 64<br />

mollis, Spirastrella, 54-55<br />

miilleri, Darwinella, 18<br />

muscarum, Ircinia, 11<br />

Mycale, 45-46; cecilia, 47-48, 84; crassissima, 51;<br />

fistulata, 48; fistulata microsigmatosa, 47, 48;<br />

isochela, 48, 50-51; /aery's, vi, 7, 46-47, 49, 77-<br />

78, 82, VI; maunakea, 48; microsigmatosa, 7,<br />

47-48, 78, 80-82, 84; parishii, 48, 50; pecrtnicola,<br />

48, 50-51; phyllophila, 48; senegalensis,<br />

47-48'<br />

Mycalidae, v, 1, 5, 7, 45<br />

Mycalinae, 45<br />

Myxilla isodictyalis, 38-39<br />

neens, Adocia, 29<br />

Ne<strong>of</strong>ibularia, 1, 6, 22-23, 84; masja, 6, 23, 79, 81<br />

nigrescens, Tedania, 37<br />

nigricans, Papillina, 61<br />

nolitangere, Fibulia, 23<br />

nucula, Chondrilla, 7, 74-75, 79, 81, 84<br />

Oligoceras, 11; collectrix, 11; collectrix hemorrhages,<br />

11; hemorrhages, vi, 6, 11, 79, 81, I;<br />

spongeliformis, 11<br />

Ophlitaspongia membranacea, 44<br />

Ophlitaspongiidae, 44-45<br />

orientalis, Cliona, 57<br />

ornata, Displastrella, 59<br />

oras, Ircinia, 10<br />

Osculina polystomella, 61<br />

Pachychalina, areolata, 25; aurantiaca, 26; carbonaria,<br />

26-27; millepora, 26; rubens, 18-19<br />

pallescens, Spongelia, 15-16<br />

pallida, Callyspongia, vi, 1, 6, 36-37, 55, 83-84,<br />

III<br />

Pandaros, 45; acanthifolium, 45<br />

panicea, Halichondria, 53<br />

papillaris, Callyspongia, 32; Spongia, 32<br />

Papillina, arcuata, 56-57; cribrosa, 57; nigricans,<br />

61<br />

papyracea, Geodia, vi, 1, 7, 71-72, 72, 78, 83, 85,<br />

VIII; Siphonochalina, 34-35<br />

parishii, Esperella, 48, 50; Esperia, 48, 50; Myca/e,<br />

48, 50; Raphiodesma, 48, 50; Zygomycale,<br />

vi, 7,48-51,77,81-82, 84, V<br />

parthena, Microciona, 42<br />

Patuloscula, plicifera, 34; procumbens, 32


pectinicola, Mycale, 48, 50-51<br />

Pellina, 27, 29-30; carbonaria, 26-27, 30; coela,<br />

30; coeliformis, vi, 1, 6, 29-30, 29, 78-79, 83-84,<br />

III; semitubulosa, 30<br />

Phloeodictyon carbonaria, 26<br />

phyllodes, Chondrilla, 54-55; Chondrillina, 54<br />

phyllophila, Mycale, 48<br />

Placospongia, 62; carinata, vi, 7, 62-64, 77, 79,<br />

81, 84, VII; intermedia, 62, 64; melobesioides,<br />

63-64; m/xta, 62, 64; sp., 62, 64<br />

Placospongiidae, v, 7, 62<br />

Plakinidae, v, 8, 76<br />

Plakortis, 4, 76; simplex, 76; zyggompha, 8, 76,<br />

79,81<br />

plicifera, Callyspongia, vi, 4, 6, 32, 34-36, 79, 82,<br />

IV; Patuloscula, 34; Spinosella, 34-35; Spongia,<br />

34-35; Tu&a, 34-35<br />

Plumocolumella, 23<br />

plumosa, Esperella, 48, 50; Esperia, 48, 50<br />

polystomella, Osculina, 61<br />

Poterion atlantica, 57-58<br />

praetexta, Aplysina, 13<br />

procumbens, Patuloscula, 32<br />

Prosuberites, 56<br />

proxima, Thalysias, 26-27<br />

pulvinata, Spirastrella, 57<br />

pulvinatus, Hymeniacidon, 57-58<br />

purpurea, Spirastrella, 54-55, 57<br />

purpureum, Alcyonium, 54<br />

Pyxitis gibberosa, 69-70<br />

quadripartitum, Corticium, 77<br />

ramosa, Fibularia, 22; Gelliodes, 22; Ircinia, 9<br />

raphidifera, Fibulia, 23<br />

Raphiodesma parishii, 48, 50<br />

rarispinosa, Microciona, vi, 1, 7, 42-44, 43, 77-78,<br />

83, 85<br />

Reniera digitata, 37-38<br />

reptans, Desmacidon, 21-22<br />

reticulata, Geodia, 69-70<br />

rhizophyta, Cinachyra, 4, 6<br />

rhoda, Ulosa, 51-52<br />

ridleyi, Esperella, 48, 50<br />

Rodsa, 76; zyggompha, 76<br />

rosacea, Darwinella, 1, 6, 17-18, 77-79, 82, 84<br />

rubens, Chalina, 18-19; Haliclona, 5, 6, 18-19,<br />

79, 83; Pachychalina, 18-19; Spongia, 18-19<br />

rubiginosa, Cacochalina, 19<br />

Sarcotragus, 10<br />

scrobiculata, Callyspongia, 32; Spinosella, 34-35;<br />

Spongia, 34-35; Tu&a, 32, 34-35<br />

sebae, Luff aria, 13<br />

semitubulosa, Pellina, 30<br />

senegalensis, Mycale, 47-48<br />

seychellensis, Alemo, 65-66; Donatia, 65-66;<br />

7>%a, 7, 65-66, 67, 78-79, 81-82, 84<br />

Sigmadocia, 30-31; caerulea, vi, 1, 5-6, 26, 30-31,<br />

30, 78-79, 81, 84, III<br />

similis, Lissodendoryx, 38-40; Microciona, 43-44<br />

INDEX 93<br />

simplex, Plakortis, 76<br />

Siphonochalina, papyracea, 34-35; stolonifera, 34<br />

saroria, Spinosella, 33; Tuba, 33<br />

souriei, Acanthacarnus, 7, 40, 79, 83-84<br />

spengelii, Aplysina, 4<br />

Spheciospongia, 57-58; vesparia, 4, 7, 57-58, 78,<br />

80<br />

spinosa, Microciona, 43<br />

Spinosella, maxima, 4, 34-35; plicifera, 34-35;<br />

scrobiculata, 34-35; sororia, 33<br />

Spirastrella, 54, 58; andrewsii, 4, 57-58; coccinea,<br />

7, 54-55, 79, 81, 83-84; coronaria, 56; cunctatrix,<br />

54-55; cunctatrix porcata, 55; cunctatrix<br />

robusta, 55; decumbens, 54-55; mollis, 54-55;<br />

pulvinata, 57; purpurea, 54-55, 57<br />

Spirastrellidae, v, 7, 54<br />

Spongelia, elastica lobata, 15-16; elastica lobosa,<br />

15-16; elastica massa, 15-16; fragilis, 14-16;<br />

incrustans, 15-16; pallescens, 15-16<br />

spongeliformis, Cacospongia, 11; Oligoceras, 11<br />

Spongia, angulosa, 51; carbonaria, 26-27; cty-<br />

5onz, 57-58; fasciculata, 8-9; fistularis, 12;<br />

fragilis, 14-15; linteiformis, 10; papillaris, 32;<br />

plicifera, 34-35; rubens 18-19; scrobiculata,<br />

34-35; strobilina, 10; tubaeformis, 12; vaginal,<br />

32-33<br />

spongia, Ulosa, 51-52<br />

Spongiidae, v, 6, 8<br />

stolonifera, Siphonochalina, 34<br />

strobilina, Hircinia, 10-11; Ircinia, 6, 9, 10-11,<br />

77-81, 84; Ircinia, irregularis, 10-11; Spongia,<br />

10<br />

Strongylacidon intermedia, 51<br />

Suberites, angulospiculatus, 4, 6; coronarius, 4,<br />

56; tuberculosus 56-57; undulatus, 62<br />

Suberitidae, v, 7, 59<br />

subulata, Cliona, 61-62<br />

Synops (?) media, 69<br />

Tedania, 37-38; anhelans, 37-38; orwcei, 37-38;<br />

digitata, 37-38; digitata bermudensis, 37-38;<br />

ignw, vi, 5, 7, 37-38, 39, 77-79; 83-84, V;<br />

ignw pacifica, 37; leptoderma, 38-39; nigrescent,<br />

37<br />

Tedaniidae, v, 7, 37<br />

tenuissima, Verongia, 12<br />

Terpios, 59; ztteftt, vi, 5, 7, 59-60, 78-79, 81-84,<br />

VII<br />

Tethya, 65-66, 81; actinia, 7, 66-67, 79, 84; dip/oderma,<br />

66-67; ingalli, 65-67; lyncurium, 65;<br />

maza, 7, 65-66, 67-68, seychellensis, 7, 65-66,<br />

67, 78-79, 81-82, 84; sp., 7, 67, 68, 79<br />

Tethyidae, v, 7, 65<br />

tetralophum, Corticium, vi, 1, 8, 77, 77, 79, 83,<br />

85<br />

Thalyseurypon, 44-45; carteri, 45; conulosa, vi,<br />

1, 7, 44-45, 44, 79, 82, 85, VI; foliacea, 45;<br />

vasiformis, 45<br />

Thalysias, carbonaria, 26-27; coccinea, 54-55;<br />

fgnt5, 37-38; proxima, 26-27; varians, 55-57;<br />

vesparia, 57-58<br />

transiens, Erylus, 74


94<br />

Tuba, plicifera, 34-35; scrobiculata, 32, 34-35;<br />

sororia, 33; vaginalis, 32-33<br />

tubaeformis, Spongia, 12<br />

tuberculosa, Geodia, 69-70<br />

tuberculosus, Suberites, 56-57<br />

tubifera, Adocia, 28<br />

tumultosa, Geodia, 69-70<br />

£/Z


PLATES


PLATE I<br />

Fig. 1. Verongia fistularis (Pallas) .......................................... p. 12<br />

Ojff Maiden Cay, Jamaica. YPM No. 5149. x0.7.<br />

Figs. 2, 3. Haliclona erina de Laubenfels ................................... p. 19<br />

Port Royal Lagoon, Jamaica, YPM No. 5166. x0.75.<br />

Fig. 4. Haliclona doria de Laubenfels ....................................... p. 18<br />

Port Royal Lagoon, Jamaica. YPM No. 5159.- x0.9.<br />

Fig. 5. Oligoceras hemorrhages de Laubenfels ................................ p. 11<br />

Port Royal, Jamaica. YPM No. 5144. X 1.1.<br />

96


4*<br />

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•<br />

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PLATE II<br />

Fig. 1. Haliclona hogarthi n. sp. ........................................... p. <strong>20</strong><br />

Port Royal, Jamaica. Holotype. YPM No. 5033. x0.8.<br />

Fig. 2. Adocia implexiformis n. sp. .......................................... p. 27<br />

Port Royal, Jamaica. Holotype. YPM No. 5034. x0.9.<br />

Fig. 3. Gelliodes areolata (Wilson) .......................................... p. 25<br />

Port Royal, Jamaica. YPM No. 5186. x0.6.<br />

Fig. 4. Desmapsamma anchomta (Carter) p. 21<br />

Port Royal, Jamaica. YPM No. 5174. xO.93.<br />

§7


PLATE III<br />

Fig. 1. PelUna coeliformis n. sp. (growing on Geodia gibberosa Lamarck) ........ p<br />

Port Royal, Jamaica. Holotype. YPM No. 5086. x0.9.<br />

Figs. 2, 8. Adocia carbonaria (Lamarck) ...................................... p<br />

Dunn's River, Jamaica. YPM No. 5191. xO.75.<br />

Fig. 4. Sigmadocia caerulea n. sp. ........................................... p<br />

Port Royal, Jamaica. Holotype. YPM No. 5037, X LI.<br />

Fig. 5. Callyspongia pallida n. sp. ........................................... p<br />

Port Royal, Jamaica. Holotype. YPM No. 5088. _x0.8.<br />

98


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• > •<br />

\^<br />

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, R-<br />

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••XV


PLATE IV<br />

Fig. 1. Callyspongia plicifera (Lamarck) p. 34<br />

Off Maiden Cay, Jamaica. YPM No. 5215. xO.55.<br />

Fig. 2. Callyspongia fallax Duchassaing and Michelotti ....................... p. 31<br />

Off Maiden Cay, Jamaica. YPM No. 5<strong>20</strong>5. xO.55.<br />

ft


PLATE ¥<br />

Fig. 1. HaMchondria mogniconulosa n. sp. ................................... p. 58<br />

Port Royal, Jamaica. Holotype. YPM No. 5089. x0.75.<br />

Fig. 2. HaMchondria melanadocia de Laubenfels .............................. p. 52<br />

Port Royal, Jamaica. YPM No. 5219. x0.85.<br />

Fig. 8. Zygomycale parishii (Bowerbank). .................................... p. 48<br />

Port Royal, Jamaica. YPM No. 5258. x0.8.<br />

Fig. 4, Tedania ignis (Duchassaing and Michelotti) .......................... p. 87<br />

Port Royal, Jamaica. YPM No. 5229. x0.85.<br />

100


->-" .*^-^; .'-fcL<br />

•"i-S; - "- i-Vi-" . • v9r£K* •'•'<br />

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~ArW$^ £-<br />

^JIL


PLATE ¥1<br />

Fig. 1. Thalyseurypon conulosa n. sp. ....................................... p. 44<br />

Off Maiden Cay, Jamaica. Holotype. YPM No. 5042. X1.<br />

Fig. 2. Mycale laevis (Carter) ............................................... p. 46<br />

Port Royal, Jamaica. YPM No. 5240. xl.l.<br />

101


PLATE VII<br />

Fig. 1. Placospongia carinata (Bowerbank) .................................. p<br />

Port Royal, Jamaica. YPM No. 5279. x L<br />

Fig. -.2. Diplastrella megastellata n. sp. ................... v...-.....'.'...'..'.'.....'..• p<br />

Off Maiden Cay, Jamaica. Holotype. YPM No. 5044. x 0.9..<br />

Fig. S. Terpios zeteki (de Laubenfels) ........................................ p<br />

Port Royal, Jamaica. YPM. No. 5270. X1.'<br />

Figs. 4, 5. Anthosigmella varians (Duchassaing and Michelotti). ................. p<br />

Port Royal, Jamaica. YPM No. 5262. x0.6.<br />

102


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PLATE VIII<br />

Fig. 1. Geodia papyracea n. sp. p. 71<br />

Port Royal, Jamaica. Paratype. YPM No. 5311. x 1.1.<br />

Fig. 2. Geodia papyracea n. sp. (specimen on lower right, growing on Geodia<br />

gibberosa Lamarck) ..................................................... p. 71<br />

Port Royal, Jamaica. Paratype. YPM No. 5311. x 1.1.<br />

Fig. 3. Erylus ministrongylus n. sp. ......................................... p. 72<br />

Drunkenman's Cay, Jamaica. Holotype. YPM No. 5046. x 1.6.<br />

103

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