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HARVARD UNIVERSITY

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LIBRARY

OF THE

Museum of Comparative Zoology

Peabody Museum of Natural History
Yale University
Bulletin 12

Natural History of the
Marine Sponges
of Southern New England
by
Willard D. Hartman

PEABODY MUSEUM OF NATURAL HISTORY, YALE UNIVERSITY

BULLETIN 22

Natural History of the

Marine Sponges
of Southern New England

BY

WILLARD D. HARTMAN

Peabody Museum of Natural History
and
Department of Zoology
Yale University

NEW HAVEN, CONNECTICUT
1958

Based on a dissertation submitted to fulfill in part the require-
ments for the degree of Doctor of Philosophy in Yale University.

Printed in the United States of America

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JAN 1 3.1959

Dedicated to

PROF. ALEXANDER PETRUNKEVITCH
on the occasion of his eightieth birthday,
22 December 1955

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CONTENTS

ABSTRACT Vil
PREFACE ix
I. SysTEMATIC STUDIES 1
Previous work 1

List of Demospongiae described from southern New England 3
Descriptions of species 3
Suberites ficus (Johnston) Schmidt 3

Cliona celata Grant 16

Cliona lobata Hancock 19

Cliona vastifica Hancock 7

Cliona truitti Old 22
Halichondria bowerbanki Burton 24
Microciona prolifera (Ellis and Solander) Verrill 36
Lissodendoryx tsodictyalis (Carter) ‘Topsent 41

Isodictya deichmannae (de Laubenfels) Hartman, new comb. 45

Haliclona oculata (Pallas) Grant 52

Haliclona loosanoffi Hartman, sp. nov. 62

Haliclona canaliculata Hartman, sp. nov. 73

Summary 76

II. Lire History Srup1es 78
Previous work 78
Methods 80
Results and discussion 87
Cliona celata Grant 87

Cliona vastifica Hancock 93

Haliclona loosanoffi Hartman of
Halichondria bowerbanki Burton 102
Microciona prolifera (Ellis and Solander) 104

Summary 104

vi CONTENTS

III. Saviniry TOLERANCE, COMPETITION FOR SUBSTRATE, AND DISTRIBU-

TION AMONG CLIONIDS 106
Introduction 106
Salinity tolerances of Cliona celata and Cliona vastifica 107
Methods 107
Results 107
Discussion 121
Osmoregulation 121
Temperature and salinity tolerance in Cliona celata 123

Salinity tolerance as a factor in the distribution and speciation
in the genus Cliona 124
Zoogeography of Cliona celata, Cliona vastifica, and other clionids 131
Introduction 131
Distribution in temperate regions 132
Distribution in the Mediterranean region 136
Further distribution 138
Summary of the zoogeography of the Clionidae 139
A note on the control of boring sponges 140
Summary 141
REFERENCES CITED 142
SYSTEMATIC INDEX 151
SuBJEcCT INDEX 153
PLATES At back of book

ABBREVIATIONS

AMNH—American Museum of Natural History
MCZ—Museum of Comparative Zoology, Harvard College
YPM—Peabody Museum of Natural History, Yale University

ABSTRACT

Twelve species of siliceous sponges from Long Island and Block Island sounds
are described. Among these are two new species: Haliclona loosanoffi, a gemmu-
liferous, encrusting haliclonid; and Haliclona canaliculata, a flat encrusting
species with perennial growth. Of previously described species, Neosperiopsis
deichmannae de Laubenfels is placed in the genus Isodictya; Chalina arbuscula
Verrill is regarded as a synonym of Haliclona oculata (Pallas).

Variations in spicule size correlated with temperature are described in Hali-
clona oculata, in which the variations are geographical, and in Halichondria
bowerbanki and Haliclona canaliculata, in which the variations are seasonal.

The period of larval settling is reported for five sponges. Annual, temperature-
correlated variations in the time of larval settling and gemmule formation are
described in Haliclona loosanoffi.

Experiments on the tolerance of two species of clionids to lowered salinities
indicate that Cliona vastifica Hancock has a wider range of tolerance than Cliona
celata Grant. A review of the distribution of these two species reveals that popu-
lations of the former have penetrated brackish waters in several parts of the
world where they show parallel trends in variation of spicule form. Competition
for substrate between the two species is discussed.

PREFACE

Although there is a wealth of literature on the taxonomy of marine sponges,
the classification of many groups of these animals remains in a state of confusion.
This is true largely because of a reliance upon skeletal characteristics in distin-
guishing species. For practical reasons, the museum zoologist has no alternative
but to regard skeletal characters of primary importance in identification. MeL
there are certain families and genera in which species cannot be identified cer-
tainly on the basis of skeletal characteristics alone, and in which attempts to do
so have led to an unnecessary multiplicity of species. There is some truth in
Bidder’s (1928) comment that “most of the alleged species have been made be-
cause the differences between spicules can be so closely measured and accurately
recorded.”

Within the family Haliclonidae, e.g., the genus Haliclona is a greatly confused
one containing a large number of species many of which are doubtless invalid.
The difficulty in this case is that the skeleton consists of only one type of spicule
(oxeas); the spicules are arranged in networks which vary from unispicular reticu-
lations to polyspicular tracts joined by isolated spicules. The networks and tracts
contain variable quantities of spongin as a cementing agent. Burton (1926) studied
the skeletal characteristics of a large series of encrusting haliclonids from a two
mile stretch of beach at Littlehampton, Sussex, and found that the skeleton varied
from a subisodictyal, unispicular meshwork with spongin restricted to the ends
of adjacent spicules to an isodictyal skeleton with multispicular tracts enclosed
by a delicate spongin sheath. He concluded that all the colonies represented
variations of a single species which he, at that time, called Reniera cinerea. In
the present studies a similar variability in skeletal structure has been demon-
strated in Haliclona oculata.

But restricting one’s attention to a study of variability in the skeletal fea-
tures of series of museum specimens can sometimes hide valid differences between
species. It has become apparent to the present writer that the challenge of un-
ravelling some of the confused synonymies among sponges can be met best
through a thorough study of populations in the field. After becoming acquainted
with living populations of sponges, species differences are not infrequently rec-
ognizable aesthetically (see Pantin, 1954) before they can be communicated to
others following the application of analytical laboratory methods. Thus, field
studies of the Long Island Sound species of Halichondria led me to regard it
as distinct from panicea long before anatomical differences in the dermal skele-
ton were demonstrable in the laboratory.

Life history studies, too, have aided in separating species in which spicule
structure and size fail to be useful. In studying the local populations of encrust-
ing haliclonids, I was inclined at first to classify them as the so-called cosmopolitan
species, Haliclona permollis, which I knew from literature studies only. Field
studies of the annual cycle of the Long Island Sound haliclonids revealed that
they die back in late fall and live through the winter as gemmules, a habit un-
known in permollis. During the course of a recent reéxamination of the local
haliclonids, I became aware of some specimens which looked different from the
more common gemmule-bearing species. These colonies were found to remain
alive throughout the winter without forming gemmules. They do undergo a
partial internal degeneration, however, losing their choanocytes. It is now clear
that two species of Haliclona occur sympatrically in the vicinity of New Haven,

x PREFACE

neither of them synonymous with H. permollis. These species might well have
remained undetected if sole reliance had been placed on studies of their highly
variable skeletal characters.

In instances where several categories of megascleres occur or where micro-
scleres are present, classification on the basis of skeletal characteristics is more
certain. But here again environmental factors may have an effect on the spicu-
lation, as has been shown by Jewell (1935) in her ecological studies of spongillids
and by Jérgensen’s experimental work (1944). These authors found microscleres
more sensitive than megascleres to reduced concentrations of silicon. Several
instances of variations in spicule form apparently correlated with environmental
factors are reported in the present studies. The spicules of Haliclona oculata
increase in size in colder waters; those of an encrusting haliclonid and a Hali-
chondria found in Long Island are larger in winter specimens. The occurrence
of parallel variations in spicule form in brackish water populations of Cliona
vastifica in various parts of the world suggests a correlation with low salinity.
In none of these cases, however, has experimental work been carried out as yet
to ascertain which of the spicule form variations are ecophenotypic and which
genetically determined.

Embryological and biochemical data have also proven applicable to problems
of sponge systematics. Topsent (1911) found a difference in breeding seasons and
in larval structure in the difficult species, Halichondria panicea and bowerbanki;
Tuzet (1948) has presented sound evidence for the distinctness of the calcareous
sponge genera, Leucosolenia and Clathrina, on the basis of her studies of the early
developmental stages of these forms; Lévi (1953a, 1956) has found that larval
characteristics are the best ones for separating sibling species of Halisarca.

Bergmann’s studies (1949) of the sterols of sponges are helpful in defining
the families Suberitidae and Spirastrellidae and provide a possible basis for a
reéxamination of the genus Haliclona. It is probable that biochemical studies of
the components of the organic skeletal elements in sponges will yield results of
importance to understanding the classifications and phylogeny of the Porifera as
they have among the Anthozoa (see Roche and Tixier-Durivault, 1951).

The present studies concern the systematics and some ecological relationships
of the sponges of Long Island Sound and Block Island Sound. Life history studies
of five common species and the salinity tolerances of two sympatric clionids have
been investigated with a consideration of the importance of these factors in the
distribution of the sponges concerned.

Willard D. Hartman
New Haven, Connecticut
September, 1958

ACKNOWLEDGMENTS

Many persons have helped me during the course of this work, and it is a
pleasure to express appreciation to all who have so generously provided assistance.
A special debt of gratitude is owed to Professor G. Evelyn Hutchinson who sug-
gested the problem and who has been a constant source of inspiration during
its execution. I should also like to express my sincere thanks to Dr. Daniel Merri-
man who provided working space and equipment at the Bingham Oceanographic
Laboratory and to Dr. Victor L. Loosanoff who offered working facilities and
much assistance at the Milford Biological Laboratory where most of the experi-
mental work was pursued.

Drs. E. S. Deevey, Jr. and E. F. Thompson have provided many suggestions
and criticisms in regard to the ecological work. Dr. Alexander Petrunkevitch and
Dr. Grace E. Pickford have given much advice and encouragement throughout
the course of the work. Dr. Werner Bergmann has been a constant source of
encouragement and inspiration through his general interest in marine zoology,
and has contributed many specimens. In addition many members of the staff
and students at both laboratories where the work was carried out, as well as at
the Osborn Zoological Laboratory, have been of assistance at one time or an-
other. Special thanks are due Mr. Joseph Lucash, of the Milford Biological
Laboratory, who provided valuable technical assistance in the work carried out
there.

The late Dr. Stanley C. Ball kindly granted permission to examine the exten-
sive collections of sponges at the Yale Peabody Museum, before I joined the
staff of that institution. Dr. Elisabeth Deichmann kindly lent the collection of
Woods Hole sponges from the Museum of Comparative Zoology at Harvard Col-
lege. Dr. John Armstrong permitted me to study specimens from the collections
of the American Museum of Natural History.

I am also indebted to Mr. Nathan Bowman, Mr. M. D. Burkenroad, Mr. G. R.
Forster, Dr. Claude Lévi, Dr. I. M. Newell, Dr. Max Pavans de Ceccaty, Dr. R. A.
Paynter, Jr., Prof. Odette Tuzet, Mr. H. E. Warfel, Dr. D. P. Wilson, and the
Director of the Plymouth Laboratory of the Marine Biological Association of
the United Kingdom for supplementary collections of sponges made at vari-
ous places on the Atlantic Coast of North America and in Europe.

Much of the collecting in Long Island Sound was done aboard boats of the
Rowe Oyster Company, New Haven, and the Connecticut Oyster Farms, Mil-
ford. Collecting in Block Island Sound was done aboard Capt. Ellery Thompson’s
boat, the “Eleanor.”

I am indebted to Dr. Carl O. Dunbar, Director of the Peabody Museum,
for helpful criticism in editing of the manuscript. I should also like to express
my appreciation to Mr. Frederick Nigretti, formerly preparator in invertebrate
zoology at the Peabody Museum, for technical assistance during revision of the
manuscript for publication, and to Mrs. Delores McColm for typing and editing
the manuscript, and for much helpful criticism. Finally, I am deeply grateful
to Miss Shirley P. Glaser for her great care in preparing the figures.

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NATURAL HISTORY OF THE MARINE SPONGES
OF SOUTHERN NEW ENGLAND

I. SYSTEMATIC STUDIES

PREVIOUS WORK

The material on which these studies are based comprise collections made
in rather restricted areas of Long Island Sound and Block Island Sound.
The intertidal and shallow water sponge fauna was studied along the Con-
necticut shore of Long Island Sound from Hammonasset State Park to Mil-
ford Point. Deeper water collections were largely restricted to oyster beds
ranging from Branford to Bridgeport, Connecticut, down to depths of ten
meters and never more than a mile and a half from shore. One collection
was made in Clinton Harbor aboard a lobster boat. The northern shore of
Long Island, the shore of Connecticut east of Clinton and west of Milford
Point, and the deeper central part of the Sound have not been surveyed as yet.
Collections from Block Island Sound were made on commercial fishing
grounds at depths varying from 20 to 40 meters. It is probable that this fauna
penetrates the deeper waters at the eastern end of Long Island Sound.

The above-mentioned collections were supplemented by a few specimens
from both Long Island Sound and Block Island Sound in the A. E. Verrill
collections at the Peabody Museum.

The earliest study of Long Island Sound sponges was reported by Rafines-
que (1819)! who described five species from the shores of Long Island
Sound. Rafinesque’s studies were restricted to the western end of the island,
including both the northern Sound shore (Oyster Bay) and the southern
shore (Gravesend, Sandy Hook, and Bath [?]). His descriptions are poor and
unaccompanied by illustrations. Possible synonyms of his species are:

1. Spongia albescens Raf. (whitish sponge) = Suberites ficus (Johnston)

(?)

2. Spongia ostracina Raf. (oyster sponge) = Microciona prolifera
(E. & S.)

3. Spongia cespitosa Raf. (bushy sponge) — Haliclona oculata (Pallas)

4. Spongia cladonia (cladonian sponge) — Haliclona oculata (Pallas)
(?)

5. Spongia virgata (slender sponge) = Microciona prolifera (E. & S.) (?)
Verrill (in: Verrill and Smith, 1873) included several Block Island Sound
and Long Island Sound records in a report on the invertebrates of Vineyard
Sound and adjacent regions. A list of sponges mentioned by him together

1 Rafinesque regarded sponges as “marine vegetables’ since “in all those I have seen,
in Europe and America, no perceptible motion nor sensibility was to be discerned in any
stage of their existence; and those who have acknowledged their animality, bring no
stronger proof thereof than an occasional slight shrinking under the hand, and an animal
smell, which are common to some marine plants.”

72 MARINE SPONGES

with the occurrence of each species in the waters under consideration
follows:

Grantia ciliata Fleming—Rhode Island.

Ascortis fragilis Haeckel—Common in Long Island Sound (New Haven,
Thimble Islands); Watch Hill, Rhode Island.

Microciona prolifera Verrill—Abundant in Long Island Sound.

Chalina oculata Bowerbank—Off Watch Hill, Rhode Island, 4—5 fath-
oms.

Chalina arbuscula sp. nov.—Very common in Long Island Sound;
Watch Hill, Rhode Island.

Isodictya (palmata?)—Watch Hill, Rhode Island.

Halichondria panicea Johnston—Abundant off Watch Hill, Rhode Is-
land, on algae in 4—8 fathoms.

Halichondria sp. “a’’—Watch Hill, Rhode Island.

Halichondria sp. “‘b’”—Long Island Sound near New Haven.

Halisarca sp.p—Watch Hill, Rhode Island.

Cliona sulphurea Verrill—Very abundant in Long Island Sound on
oysters and other shells.

All the species listed except Halisarca, Halichondria panicea, and Hallt-
chondria sp. “a”, have been found by the present author in the course of
collecting. ‘The synonymies of the species listed by Verrill will be considered
in the descriptive section of this paper with the exception of the calcareous
sponges which will be discussed in a forthcoming paper.

Verrill (1875) described a “curious, very slender and delicate, bipin-
nately branched species from the piles both at Greenport and Noank.”
Specimens of this sponge have not been found in the Peabody Museum and
in the absence of data on its skeletal characteristics, one can only guess as to
its identity. It is possible that Verrill was referring to colonies of Haliclona
loosanoffi, a species described as new in the present paper. Verrill goes on
to state that “several sponges, new to the fauna [eastern Long Island Sound]
were also obtained and were studied by Prof. Hyatt.” Unfortunately Hyatt
did not publish his studies of these specimens and apparently failed to return
them to the Peabody Museum.

Fragments of white Italian marble from a cargo wrecked off Long Island
in 1871 were described by Verrill (1878) as being riddled by the boring
sponge, Cliona celata. ‘The cavities penetrated to a depth of one or two
inches.

Old (1941) listed four species of clionids from Long Island Sound:
Cliona celata, C. lobata, C. vastifica, and C. truitti. The first three species
have been collected on the Connecticut shores of Long Island Sound, but
the last-named species has not been found.

SYSTEMATIC STUDIES 3

LIST OF DEMOSPONGIAE DESCRIBED FROM LONG ISLAND SOUND
AND BLOCK ISLAND SOUND

CLAss DEMOSPONGIAE

ORDER HADROMERINA
Family Suberitidae
Suberttes ficus (Johnston) Schmidt
Family Clionidae
Cliona celata Grant
Cliona lobata Hancock

Cliona vastifica Hancock
Cliona truttts Old

ORDER HALICHONDRINA
Family Halichondriidae
Halichondria bowerbanki Burton

ORDER POECILOSCLERINA
Family Desmacidonidae
Isodictya deichmannae (de Laubenfels) new comb.
Family Microcionidae
Microciona prolifera (Ellis and Solander) Verrill
Family Myxillidae
Lissodendoryx tsodictyalis (Carter) ‘Topsent

ORDER HAPLOSCLERINA
Family Haliclonidae
Haliclona oculata (Pallas) Grant
Haliclona loosanoffi sp. nov.
Haliclona canaliculata sp. nov.

ORDER DENDROCERATIDA
Family Halisarcidae
Haltsarca sp. (description not included herein)

DESCRIPTIONS OF SPECIES
OrDER HADROMERINA

FAMILY SUBERITIDAE Schmidt
Suberites ficus (Johnston, 1842) Schmidt, 1870

4 MARINE SPONGES

SYNONYMY:

? Alcyonium ficus Pallas, 1766, p. 356 [partim]

? Spongia ficiformis Poiret. Lamouroux, 1824, p. 349

? Suberites ficus, Nardo, 1833, p. 523 [partim?]

Halichondria virgultosa Johnston, 1842, p. 137

Halichondria suberea Johnston, 1842, p. 139

Halichondria ficus Johnston, 1842, p. 144

Halina suberea, Bowerbank, 1861, p. 235

Hymeniacidon subereum, Bowerbank, 1863, p. 1111

Hymeniacidon ficus, Bowerbank, 1864, p. 244

Hymeniacidon virgultosa,®? Bowerbank, 1866, p. 193

Halichondria farinaria Bowerbank, 1866, p. 269

Chalina ficus, Bowerbank, 1866, p. 270 (Misprint?)

Suberites farinaria, Schmidt, 1866, p. 16

Ficulina ficus, Gray, 1867, p. 523

? Suberites liitkent Schmidt, 1870, p. 47

Suberites ficus, Schmidt, 1870, p. 76

Suberites compacta Verrill, in: Verrill and Smith, 1873, p. 744 (Non
Alcyonium compactum Lamarck, 1815, p. 166; Halichondria compacta
Lieberkiihn, 1859, p. 520)

Suberites montalbidus Carter, 1880, p. 256

? Suberites montiniger Carter, 1880, p. 256

Suberites virgultosa, Vosmaer, 1882, p. 32

Suberites suberea, Topsent, 1887, p. 150

Suberites latus Lambe, 1893, p. 71

Suberites farinarius, Hanitsch, 1894, p. 179

Suberites placenta ‘Thiele, 1898, p. 39

DescripTIon: Block Island Sound specimens of Suberites ficus (Pl. 1, fig. 5)
grow attached to dead lamellibranch shells (e.g., Arctica islandica, Volsella modi-
olus, and Venericardia borealis) or rocks. In many cases the sponges attach to the
shells of living specimens of Venericardia borealis which remain alive although
largely enclosed in the bases of sponge colonies reaching heights of 30 cm. or
more. Miiller (1914) noted a similar relationship of ficus with Astarte in the
Barents Sea. In form the Block Island Sound colonies are similar to Miiller’s
Barents Sea specimens; they grow out from the substratum to become flat, lobate
colonies which reach considerably larger sizes than those reported by Miiller,
however. As a general rule, these colonies probably lie flat on the sea bottom,
rather than grow vertically. In several cases shells have been found imbedded in
the distal ends of the colonies, suggesting that the colonies were lying flat and
had come in contact with shells on the sea bottom. Long, attenuated specimens
such as Bowerbank (1874) figures (Pl. 35, fig. 3, as Hymeniacidon virgultosa) have
not been found in this region, nor have the “swberea” types which grow around
shells occupied by hermit crabs. The largest Block Island Sound specimen col-
lected measures 36 x 16 x 3 cm. in length, width, and thickness. Typical speci-
mens measure 28 x 9 x 2.5 cm. This sponge is very common on the fishing
grounds of Block Island Sound, often filling the nets of hopeful fishermen who
refer to it contemptuously as “elephant dung.”

2It is of interest to note that in 1863 (p. 1129) Bowerbank regarded Johnston’s Hali-
chondria virgultosa as a synonym of H. ficus.

SYSTEMATIC STUDIES 5

In consistency the colonies are quite compressible and elastic. In this respect
they differ noticeably from colonies of Suberites domunculus in my possession,
the latter being firm and incompressible. Topsent (1900) pointed out this dif-
ference between the two species and attributed the greater firmness of domuncu-
lus to the smaller diameters of the aquiferous canals. My own data fail to bear
out Topsent’s assertion. The canals in Block Island Sound specimens of ficus
measure from 300 to 800u in diameter; in specimens from Roscoff, France, they
vary up to 600. On the other hand, the canals of a colony of domunculus from
Banyuls-sur-mer, France, range from 1.0 to 1.8 mm. in diameter. It seems rather
to be the greater abundance of canals in ficus which accounts for the difference
in consistency. Cut surfaces of colonies of ficus are illustrated by Topsent (1900)
and Miiller (1914); of domunculus, by Celesia (1893), Lendenfeld (1898), and
Topsent (1900). These illustrations confirm the more porous nature of the interior
of ficus as compared with domunculus. The larger quantity of spongin which
cements together the spicules in domunculus probably also adds to the firmness
of the colonies.

The dermis of Block Island Sound specimens of ficus is quite uniformly per-
forated by pores which are outlined by a network of ridges made up of numerous
microscleres packed together. In some specimens occasional tufts of megascleres
reach the surface and reinforce the microsclere spiculation there. In specimens
of ficus from Roscoff, microsceleres are less abundant and projecting tufts of
megascleres more frequent in the dermis. Indeed, such specimens appear to be
transitional from the condition in Block Island Sound specimens of ficus to speci-
mens of domunculus from the Mediterranean, where the dermis is pierced ex-
clusively by tufts of megascleres surrounding the dermal pores. Oscular and pore
sizes for specimens of ficus and domunculus are given in Table 1.

The pores of specimens of ficus which I have examined are elongate and ellip-
tical in outline, the diameters of the minor axes being about half that of the
major axes. The pores of specimens of domunculus in my possession tend to be
smaller and rounder than those of ficus. Lendenfeld (1898) found the reverse to
be true in regard to ostial sizes of ficus and domunculus. The pore and oscular
measurements probably have little significance, however, when made on pre-
served specimens. One to five oscules may occur per specimen in both species.
Small slit-like fissures, presumably formed by amphipods, are common on the
surface of some specimens of both species which I have studied (see Vosmaer,
1933).

The flagellated chambers in the two species are similar in shape (spherical or
ellipsoidal) and dimensions (see Table 2).

Small, aspiculous gemmules are present in all specimens of both species
examined by the writer. These are apparently present throughout the year, de-
veloping in contact with the substratum on which the sponge is growing. They
were noted by Topsent (1900) as well.

Spicule sizes and the proportions of the several megasclere categories present
in each case are given in Tables 3 and 4 (see also figs. 1 and 2).

Worthy of mention is the nature of the spicules present in the layer lining the
hermit crab burrow in a colony of domunculus from Banyuls-sur-mer. These
spicules are thicker than those in the rest of the sponge; included among them is
a high proportion of strongyles and tylostyles with the distal end rounded instead
of pointed (fig. 3). It is also of interest to point out that the ectosome has few
oxeas and more tylostyles than the endosome in this specimen.

SPECIES

S. ficus

S. ficus

S. ficus

S. ficus

S. ficus

S. ficus

S. ficus

S. domunculus

S. domunculus

S. domunculus

MARINE SPONGES

TABLE} 1
PORE AND OSCULE SIZES OF SUBERITES FICUS AND DOMUNCULUS

LOCALITY AND AUTHOR

Block Isl. Sound
YPM #760
(Hartman)

Block Isl. Sound
YPM #774
(Hartman)

Block Isl. Sound
YPM #779
(Hartman)

Block Isl. Sound
YPM #2119
(Hartman)

Roscoff, France
YPM #2110
(Hartman)

Adriatic Sea
(Lendenfeld, 1898)

France
(Topsent, 1900)

Naples
YPM #2029
(Hartman)

Banyuls—sur—mer
YPM #2030
(Hartman)

Adriatic Sea
(Lendenfeld, 1898)

RANGE OF PORE SIZES

86 x 74u to 45 x 25y

90 x 60u

74 x 41 to 57 x 33y
111 x 41 to 74x 37p

62 x 32 to 45 x 33yu

30 to 14u

30 to 10u

57 x 45 to 41 x 41

62 x 53u to 50 x 32u

70 to 40u

OSCULAR SIZES

2.0 mm.

2.0 mm.

closed

1.0 to 1.6 mm.

1.0 to 1.5 mm.

3.0 to 20.0 mm.

0.5 mm.

1.5 to 3.0 mm.

3.0 to 6.0 mm.

Spongin fibers are of regular occurrence in both species, domunculus and
ficus, but are more extensively developed in the former. In a specimen from Ban-
yuls-sur-mer continuous fibers enclosing tylostyles, and measuring up to 90u
across, run from the interior to the surface. In specimens of ficus from Roscoff,
small patches of spongin (up to 20u across) are found in places in the endosome
surrounding clumps of megascleres. In Block Island Sound specimens of ficus,
spongin is even rarer in occurrence.

Discussion: The tangled synonymies of Suberites ficus and Suberites domuncu-
lus have been exhaustively tabulated by Vosmaer (1933) and critically reviewed

SYSTEMATIC STUDIES iv

TABLE: 2
FLAGELLATED CHAMBER DIMENSIONS IN SUBERITES

SPECIES LOCALITY AND AUTHOR RANGE OF SIZES

S. ficus Block Isl. Sound 21 x 21p to 25 x 2ip
(YPM #2119, Hartman)

S. ficus Roscoff, France 21 x 23u to 25 x 29u
(YPM #2110, Hartman)

S. ficus Adriatic Sea 23-25
(Lendenfeld, 1898)

S. ficus France 25u
(Topsent, 1900)

S. domunculus Banyuls-sur—mer, France 25 x 25u to 29 x 33u
(YPM #2030, Hartman)

S. domunculus Adriatic Sea 25u
(Lendenfeld, 1898)

by Burton (1953). Both authors conclude that the two species are synonymous.
On the other hand, Lendenfeld (1898), Topsent (1900), Arndt (1935) and de
Laubenfels (1949), among others, have regarded them as separate species. Since
neither position is unequivocal at this time, it seems worth-while to review the
evidence once again. For the sake of discussion, it will be assumed that two species
are involved.

The spicules of Suberites domunculus (fig. 2) consist entirely of megascleres,
including tylostyles, styles, oxeas, and intermediates between these. There is gen-
erally a high proportion of oxeas present; sometimes these outnumber the other
categories. Microscleres are absent. In Suberites ficus (fig. 1), however, both megas-
cleres and microscleres occur. The megascleres are chiefly tylostyles and styles,
with some specimens containing a few oxeas, but the latter are never very abun-
dant. In addition, there are varying numbers of centrotylote microscleres, rang-
ing in shape from microstrongyles to microstyles and microxeas. Occasionally the
central swelling is absent. In any one individual both smooth and microspined
centrotylotes may occur, the proportions varying from specimen to specimen.

It is the great variation which occurs in the proportions of the several cate-
gories of megascleres and in the number of microscleres present which have led
many authors to synonymize ficus and domunculus. Thus Vosmaer (1933, p. 436)
states that the “‘presence or absence [of oxeas] has no relation to the presence
or absence of centrotylote microscleres.”” Yet his own data belie this statement.
From his table of spicule sizes (1933, pp. 455-456),3 including data of other au-
thors as well as his own, we learn the following:

8 Bowerbank’s Hymeniacidon suberea, Hanitsch’s “Suberites domuncula,” and Lambe’s
Suberites montiniger and concinnus are omitted in this discussion, the first two because
of the likelihood that spicule categories were overlooked, the last two because of their
uncertain synonymy with either of the species in question.

MARINE SPONGES

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SYSTEMATIC STUDIES 9
TABLE 4
SPICULE MEASUREMENTS OF SUBERITES DOMUNCULUS
NO. OF
LOCALITY SPICULES TYLOSTYLES STYLES OXEAS
MEASURED
Naples 100 119-209-295 168-232-303 98-259-320y
YPM #2029B x 2.5-3.7-4. 1p x 2.5-4.0-4.9u x 2.5-4.0-5.3u
28% 45% 27%
Naples 100 107-214-287 180-238-279 205-284-353
YPM #2029C Nat AU BBO 9 A a Yn) x 2.5-3.9-6.2p
50% 26% 24%
Naples 100 90-199-324u 115-236-365 115-285-390pu
YPM #2029A x 3.7-4.6-6.2u x 3.7-4.4-5.3u x 3.3-4.3-+5.7u
23% 29% 48%
Naples 50 98-258-398u x 3.7-5.6-7.8u
Osborn Zool. Lab. Colls. 59% 22% 19%
Categories not separated in measurements
100 Dermis:
90-182-320u 99-217-349u 279-348-390u
x 3.3-5.0-7.4u x 4.1-5.2-7.8y x 4.1-5.0-7.0u
44% 50% 6%
100 Endosome:
144—280-332u 144-303-373u 275-364-418u
x 4.1-5.6-7.8u x 4.5-5.9-8.2u x 4.1-5.5-7.8y
24% 47% 29%
Banyuls-sur-mer, France —_—
YPM #2030 100 Spicules lining crab burrow:
221-268-320u 234-269-303 271 x 7.4y
x 4.5-7.5-9.Ou x 7.4-8.2-9.8u
27% 8% 1%

Other categories:
Tylostyles with rounded distal ends:
144-22]-275u x 6.2-8 .0-9.4u

31%

Strongyles:
148-214-312u x 4.5-7.8-9.8u

33%

10 MARINE SPONGES

A
B
SOy Ea ee}

FicurE 1. Spicules of Suberites ficus. A. Styles and tylostyles, some of latter with

subterminal heads. B. Centrotylote microxeas, microstrongyles and microstyles.
Block Island Sound. YPM +779.

\

Figure 2. Spicules of Suberites domunculus: oxeas, styles, tylostyles. Banyuls-sur-
mer, France. YPM 72030.

Ficure 3. Peculiar strongyles and tylostyles of Suberites domunculus found in the

basal layer in contact with the enclosed hermit crab. Banyuls-sur-mer, France.
YPM #2030.

SYSTEMATIC STUDIES ll

Specimens reported with oxeas and no microscleres—9
Specimens reported with microscleres and no oxeas—15
Specimens reported with both microscleres and oxeas—3
Specimens reported with neither microscleres nor oxeas—2

It is apparent that the oxeas and microscleres are mutually exclusive in occur-
rence, as a general rule. ‘The three instances in which both microscleres and oxeas
occur can be considered specimens of S. ficus with a few oxeas; the two cases of
individuals lacking both oxeas and microscleres can be interpreted as specimens
of S. domunculus without oxeas. Vosmaer’s own records (the last eight specimens
in his table) include seven specimens with oxeas varying in number from mod-
erately frequent to numerous and with no microscleres. In the remaining speci-
men, microscleres are present and oxeas are described as “very rare.”

Burton (1932) states that the presence or absence of microscleres is valueless
as a taxonomic character for distinguishing ficus from domunculus because the
microscleres in ficus are often scarce and hence difficult to find. However, in my
experience, the simple expedient of centrifuging a suspension of spicules freed
by nitric acid from a cubic centimeter of a specimen has proved a satisfactory
means of finding rare categories of microscleres. It is certainly true that centro-
tylotes are uncommon in occasional specimens of ficus. It is even possible that
some specimens of ficus lack microscleres entirely, but I know of no cases reported
in the literature where proper precautions were taken to exclude the possibility
that microscleres were overlooked.

It seems highly probable that northern records of S. domunculus reported in
the literature are always misidentifications of S. ficus, in which the scarcity of
microscleres has rendered them undetected. Several instances in which subse-
quent examinations of specimens of S. ficus have revealed the presence of micro-
scleres which had been overlooked previously may be cited. Topsent (1900) has
pointed out, for example, that Carter found centrotylote microstrongyles upon
reéxamining the specimens which Johnston (1842) and Bowerbank (1866) had
described as Halichondria suberea and Hymeniacidon subereum, respectively, a
species lacking microscleres according to these authors. Schmidt (1870) synony-
mized suberea with Suberites domunculus, also having overlooked the micro-
scleres. Hanitsch (1890), who accepted Schmidt’s synonymy of swberea with do-
munculus, found no microscleres in specimens from the Liverpool region. Later
(1891), however, he did find them in his specimens of “suberea’” but he con-
tinued to call the species domunculus and to distinguish it from ficus without
mentioning his reasons for this. Lambe (1894) pointed out that he had overlooked
the microscleres in some of his specimens of Suberites latus, which he had de-
scribed previously (1892) as a new species. De Laubenfels (1949) identified a speci-
men from Woods Hole as Suberites domunculus on the basis of the absence of
microscleres, but a reéxamination of his specimen (MCZ, +6509) by the present
author has revealed that microscleres are not uncommon in it. Records of Suber-
ites domunculus (as defined here) from the Atlantic coast of France, Great Britain
and the North Sea must all remain in doubt until the absence of microscleres has
been proven by careful reéxamination.

In addition to the differences in spicule types between the two species in ques-
tion, the abundant literature on these sponges reveals distinctions in form and
habitat. These characters overlap in part in the two species, a fact which has aug-
mented the confusion.

12 MARINE SPONGES

Suberites domunculus almost always grows around a gastropod shell inhabited
by a hermit crab and has a massive, rounded form. Occasionally this species grows
on the backs of dromiid crabs. Suberites ficus is more variable in form and habi-
tat. Some specimens (“var. swbereus”) grow on shells occupied by hermit crabs and
develop into massive, rounded colonies (Miller, 1914, e.g., found this form to be
the most common one in the North Sea); others grow on shells of living or dead
gastropods and lamellibranchs or on rocks and assume a figlike shape (which gives
the sponge its specific name) or form elongate, flattened colonies which are some-
times lobose (Miiller, 1914, found the latter shape common in the Barents Sea);
still others (‘‘var. farinarius’’) have an elongate, cylindrical form and grow at-
tached to shells or rocks.

Turning now to the geographical distribution of the two species we find that
S. ficus is a wide ranging species, having been reported from the Mediterranean
area and the west coast of Africa to Senegal, from the west coast of France, the
English Channel, the North Sea, the coasts of the British Isles and Norway, off
Iceland and Greenland, south along the Atlantic Coast of North America to Vir-
ginia, from the Bering Sea, the Pacific Coast of North America south to Van-
couver Island, and from the Pacific Coast of Asia to Japan.

S. domunculus, on the other hand, has a restricted range, overlapping that of
ficus, in part. S. domunculus occurs only in the Mediterranean area, including
the Black Sea, and southward along the west coast of Africa to Senegal. The
Suberites recorded by Lendenfeld (1888) from the south and east coasts of Aus-
tralia is of doubtful synonymy with domunculus. Suberites heros Schmidt (1870),
from the Antilles, has been considered a synonym of domunculus by ‘Topsent
(1900), Vosmaer (1933), Arndt (1935), and Burton (1953), but it seems likely that
Thiele (1905) is correct in regarding it as a valid species. Burton’s record of
domunculus from South Saghalin (1932) is questionable.

The information from the literature here reviewed may be interpreted in one
of two ways: (1) the two species are distinct, ficus being a wide ranging species
with domunculus occurring sympatrically with it over part of its range; (2) there
is only one species which is polymorphic in the Mediterranean area and along
the northwest coast of Africa. In my opinion, the first interpretation best fits the
data reviewed.

Suberites domunculus, then, is a species which (1) has a relatively constant ex-
ternal form, (2) invariably occurs in association with hermit crabs or dromiid
crabs, (3) lacks microscleres, (4) tends to have a high proportion of oxeas among
the megascleres, and (5) has a restricted geographic range. S. ficus, on the other
hand, (1) is a highly variable species in regard to external form and habitat, over-
lapping in part with S. domunculus in these characteristics, (2) always possesses
microscleres, although the number present may vary greatly among individuals
from a single population, (3) usually lacks oxeas, and (4) has an extensive geo-
graphic range.

It is hoped that this conclusion, based on previous work in the literature and
a study of the limited material available to me, will be checked through a fresh
attack on the problem. Some possible approaches follow: Biometric studies of
large series of specimens from populations of Suberites in the Mediterranean, the
west coast of Europe, the Arctic, and North America, would help to determine
the significance of variations in spicule categories and external shape. Cytological
studies, including chromosome counts, might prove useful, although the latter
would be difficult because of the small size of the cells. Histochemical and bio-

SYSTEMATIC STUDIES 13

chemical studies of the organic skeletal elements would be of interest. Herlant-
Meewis (1949) reported the presence of spongin, elastin fibers, and cell cords in
Suberites domunculus. Are the supplementary skeletal elements similar or differ-
ent in S. ficus? Finally, further studies of development might reveal specific dif-
ferences. Topsent (1900) described the eggs of S. ficus; Thomson (1887) described
both eggs and sperm in the same individual of S$. domunculus studied at the
Berlin Aquarium.

The possibility exists that ficus itself represents a complex of species. There
is a reasonable doubt as to whether Suberites liitkeni and S. placenta are syn-
onyms of ficus, for example, and the populations of ficus occurring south of Cape
Cod on the North American Coast (Verrill’s S. compacta) may actually represent
a distinct species, although further analysis is necessary to establish this.

NOTES ON RELATIONSHIPS AND NOMENCLATURE: Suberites ficus is often placed
in the genus Ficulina, but this seems unwarranted in view of the close similarity
of ficus to domunculus. Perhaps Ficulina is best considered as a subgenus of
Suberites (as suggested by Thiele, 1905, p. 416) to receive those species having
centrotylote microscleres, such as ficus and liitkeni (if the latter is indeed distinct
from ficus).

De Laubenfels (1936, 1949) has placed S. ficuws in the family Spirastrellidae* on
the basis of the presence of microscleres; thus he separates ficus widely from the
Suberitidae.> However, this author has placed undue emphasis on the occurrence
of microscleres and has ignored the striking similarities between ficus and domun-
culus in general morphology. Moreover, Bergmann (1949) has demonstrated that
a clear-cut difference exists in the sterols of spirastrellids and suberitids. He
found clionasterol and poriferasterol in Spheciospongia and Anthosigmella, and
cholestanol and neospongosterol in Suberites domunculus and Suberites ficus
(the latter species is reported by him as Suberites suberea [from Alaska] and
S. compacta [from Block Island Sound]). The biochemical evidence is in agree-
ment with the morphological similarities of S. ficus and S. domunculus, and it
must be concluded that the two species cannot be separated at the generic and
familial levels.

If Suberites ficus is accepted as a species distinct from S. domunculus, it is a
dificult matter to decide upon a valid name for it. Since authors prior to John-
ston (1842) failed to mention the microscopic details necessary to separate ficus
from domunculus and carnosus, it seems to me impossible to accept a name pro-
posed before 1842.

Pallas (1766) is the first author after 1758 who refers to a sponge which might
be Suberites ficus. His Alcyonium ficus actually includes both a sponge and a
compound ascidian [Synoicum pulmonaria (Ellis and Solander)], as pointed out
by Hartmeyer (1914). Pallas had apparently not seen a specimen (“mihi nunquam
visum”) of either the British sea-fig (an ascidian) or the Mediterranean sea-fig (a

*De Laubenfels (1956, 1949) named the family “Choanitidae” because Mantell (1822)
had established Alcyonium ficus Linné as the type of Choanites, a genus of fossil sponges
which have no apparent relationship to the recent sponge, Suberites ficus. Topsent (1933)
and Burton (1953) have accepted the view of Lamouroux (1824) that Linné’s Alcyonium
ficus is an ascidian, and de Laubenfels (1955) has now dropped the name Choanites with-
out giving a generic assignment to the species ficus.

* Burton (1953) states that de Laubenfels (1949) “appears to accept the identity of
Ficulina ficus with Suberites domuncula,” but in reality de Laubenfels places the two
species in different families in the paper under consideration.

14 MARINE SPONGES

sponge), and was misled by the similarity of the figures of these animals given by
earlier authors. He cites Imperato (1599) who figures a sponge called Alcyonium
tuberosum forma ficus; Bauhino and Cherlero (1651) who give a copy of Imper-
ato’s figure; Mercati (1717) who describes what may be Suberites ficus, naming it
Alcyonium quintam antiquorum®, Ray (1724) who does not refer to a sponge;
Marsilli (1725) who figures a sponge which may have been Suberites ficus, but
looks more nearly like Petrosia ficiformis (Poiret); and Ellis (1755) who clearly has
reference to the ascidian. It is entirely possible that some of Pallas’ indications do
indeed refer to Suberites ficus, but there is no way of being certain of this since
the figlike form shown in the figures of all of these authors is assumed occasionally
by Suberites carnosus and possibly also by domunculus. Pallas’ name is thus
without significance today. The same is true of Linné’s (1767) name Alcyonium
ficus; this author simply cites a few of the same authors mentioned above, namely,
Bauhino and Cherlero, Ray, Marsilli, Ellis, and Pallas. Linné’s use of ficus also
includes both an ascidian and a sponge, the latter undeterminable. Battarra
(1773) cites Imperato’s Alcyonium tuberosum Ficus forma (sic) without descrip-
tion or figure; Ellis and Solander (1786) figure a sponge called “‘the sea-fig” which
they clearly differentiate from the compound ascidian, but they do not give a sci-
entific name for it. Hartmeyer (1914) lists other references to Alcyonium ficus
which are either undeterminable or include both an ascidian and a sponge.

Poiret (1789) mentions an Alcyonium ficus which is certainly a sponge (it is
described from the Mediterranean region which is outside the range of the ascid-
ian; Poiret also mentions the presence of an oscule); although this is probably a
Suberites [he calls it “the fig-shaped sea cork,” thus anticipating the name Suber-
ites which was first used by Nardo (1833)], it is not possible to say with certainty
to which species it belongs. Poiret differentiates Alcyonium ficus from another
sponge, similar in external shape, which he calls Spongia ficiformis. This species,
now called Petrosia ficiformis (Poiret), adds further confusion to the literature,
although Nardo (1844) had recognized Lamarck’s citation of this species as dis-
tinct from Suberites ficus and had named it Reniera dura var. ficiformis. Burton
(1953) has tabulated Topsent’s (1933) views on the early nomenclatural history
of this species. Topsent points out that although Poiret’s Spongia ficiformis, and
most subsequent references to it, actually refer to Petrosia ficiformis, the citation
of this species given by Lamouroux (1824) probably refers to Suberites ficus, the
name of which should be changed to fictformis. But once again Lamouroux’s
description is open to confusion with other species of Suberites, and this name
cannot be accepted for Suberites ficus in my opinion.

Burton accepts Topsent’s view that “all references to the so-called Ficulina
ficus prior to Lamouroux (1824) are concerned with either Petrosia ficiformis
Poiret or an ascidian.” He thereby rejects the conclusions of Hartmeyer, who, in
my opinion, has given a sounder review of the early literature. There is little
doubt in my mind that some of the authors prior to Lamouroux did indeed have
reference to a sponge other than Petrosia ficiformis, and this sponge was most
probably a Suberites. But I do not feel that any of these authors has given a de-
scription which enables ficus to be distinguished from other member of the
genus.

6 Mercati gives three figures of this animal. Two have the figlike form portrayed
earlier by Imperato (1599), but only one of these is shown with a large central oscule and
may represent a Suberites. The third animal figured is lobate and lacks distinct oscules.

SYSTEMATIC STUDIES 15

Nardo (1833) established the genus Suberites to include the following species:
“Suberites typus N., Alc. domuncula Olivi, S. ficus N., Sp. ficus? auct., S. volubilis
N.” The first and last mentioned species are nomina nuda, no description being
given in the paper mentioned or in any of his subsequent works. (It is possible
that these species are described in his posthumous publications (1847) which I
have not seen.) It is of interest to note that Nardo questioned the synonymy of
his Suberites ficus with Spongia ficus auct. In 1844 Nardo clarified his concept of
Suberites (which he now spelled Suburites) ficus, describing it as a synonym of
Alcyonium ficus Olivi and Ginnani’s (1757) Alcyonio minore in forma di fico
frutto. Although Nardo described the megascleres of this species, he did not men-
tion the diagnostic microscleres.

If the presence of microscleres is accepted as a valid character for distinguish-
ing ficus from domunculus, then the type chosen should make reference to their
presence. So far as I have been able to determine from the literature, Johnston’s
(1842) descriptions of Halichondria virgultosa, H. suberea, and H. ficus are the
first unequivocal references to the sponge in question. Johnston does not men-
tion the presence of microscleres in any of these species, but their occurrence has
been affirmed in all by subsequent authors who reéxamined Johnston’s material.
The name virgultosa™ has page priority (p. 137); the presence of microscleres in
Johnston’s specimen was confirmed by Bowerbank (1866, pp. 193-195).8 The
species suberea is mentioned next (p. 139); the presence of microscleres in John-
ston’s specimen was confirmed by Carter (Hanitsch, 1891, p. 218). The name ficus
is listed last of all (p. 144); Bowerbank (1864, p. 244 and Pl. IV, fig. 95) figured
a microsclere from a specimen which he regarded as identical to those of John-
ston. The specific name virgultosa has priority on the basis of strict adherence to
the Rules of the International Commission on Zoological Nomenclature. How-
ever, the name ficus has become so well established that it seems desirable to con-
serve it, especially since this is the first name which was used with a figure of the
diagnostic microsclere (Bowerbank, 1864). Schmidt (1870) first referred Bower-
bank’s Hymeniacidon ficus [= Halichondria ficus Johnston] to the genus Suber-
ites.

The name domunculus was first used by Olivi (1792); as Burton (1953) has
noted, if both ficus and domunculus are accepted as valid species, it is impossible
to say which of the two species Olivi had at hand, since both may be found on
shells occupied by hermit crabs. Lieberkiihn (1859) was apparently the first au-
thor to describe the skeletal characteristics of domunculus as this species is under-
stood in the present paper. This author uses the name Halichondria compacta,
however, which should therefore replace domunculus.

Thus the names of both of these common sponges, which have become well
known in the literature as domunculus and ficus, would appear to be invalid.
The present writer feels that there is ample reason to conserve the commonly
used names, and the question will have to be submitted to the International
Commission on Zoological Nomenclature. It seems unwarranted to follow this
course, however, until the present uncertainty about the distinctness of the two

7 Johnston’s citation of Lamarck’s Spongia virgultosa is open to question. Topsent
(1933) was unable to find Lamarck’s specimen, which remains undeterminable.

8 Burton (1953) maintains that Johnston’s Halichondria virgultosa is not the same
species as Bowerbank’s Hymeniacidon virgultosa, but Bowerbank (1866) states, “The type
specimen of this species, described by Dr. Johnston . . . is in the possession of Mr. Bean,
of Scarborough, where I have had the pleasure of seeing it.”

16 MARINE SPONGES

species is clarified. In the meantime, retention of the commonly used specific
names would seem to be the best procedure to follow.

DISTRIBUTION OF Suberites ficus IN NoRTH AMERICA: Atlantic Coast of
North America, Hudson Bay to Virginia; Canadian Arctic; Pacific Coast of
North America, Bering Sea to Vancouver Island.

Vancouver Island and mainland of British Columbia, 12 to 45 meters (as S.
latus, Lambe, 1892, p. 72); Bering Sea and North Pacific Ocean (as S. suberea,
Lambe, 1894, p. 127; 1900a, p. 161); Unalaska Island (as S. montalbidus, Lambe,
1894, p. 128); Bernard Harbor and Stapylton Bay, North West Territory, 6 to
28 meters (as Ficulina ficus, Dendy and Frederick, 1924, p. 6); Hudson Bay, 35 to
55 meters (as S. montalbidus, Lambe, 1900b, p. 24); Gulf of St. Lawrence (fide
Procter, 1933, p. 108); Sable Island, N.S. (Lambe, 1896, p. 193); Mt. Desert Island
Region, Maine, 52 meters (as S. montalibidus, Procter, 1933, p. 108); Cape Cod,
Mass., 28 meters (as S. compactus, Verrill, 1880, p. 232); off Martha’s Vineyard,
Mass., 18 meters (as S. compactus, Verrill, in: Verrill and Smith, 1873, p. 744);
Nantucket (as S. compactus, Verrill, ibid., and 1880, p. 232; Sumner, Osburn,
and Cole, 1913, p. 558); Woods Hole, Mass. (as Choanites ficus and Suberites
domunculus, de Laubenfels, 1949, pp. 19, 20); Block Island Sound, 20 to 40
meters (Hartman); eastern shore of Virginia (as S. compactus, Verrill, in: Verrill
and Smith, 1873, p. 744).

FAMILY CLIONIDAE Gray
Cliona celata Grant, 1826b

SynonyMy: See Topsent (1900, pp. 32-34) and Vosmaer (1933, pp. 349-383).
As usual Vosmaer has obscured relationships within the Clionidae by excessive
lumping. Such species as Cliona lobata Hancock, Cliona viridis (Schmidt) Fischer,
and Cliona schmidti (Ridley) Topsent are unquestionably valid species although
Vosmaer regards them as synonyms of Cliona celata. Cliona caribboea Carter
may be a synonym of celata (see de Laubenfels, 1936, p. 155), although its pattern
of growth in the y-stage is certainly different from that of celata.

Discuss1on: Cliona celata is a very abundant sponge in Long Island Sound
where it is considered a pest by oyster farmers. All three stages named by Vos-
maer (a-, f-, and y-stages) are found, the a-stage, (Pl. 1, figs. 1, 2) being most com-
mon, living in galleries which it excavates in the shells of oysters and other mol-
luscs (see Part II). The f-stage (Pl. 1, fig. 3) is a transitory one passed through as
the sponge overgrows the calcareous substratum on which it first settles. The free-
living y-stage (PI. 1, fig. 4) is especially common on abandoned, undisturbed oyster
beds, as would be expected, and it is not infrequently found at mean low water
level growing on rocks. Intertidal colonies remain alive and active through the
winter. The y-stage colonies usually have no trace of the original calcareous sub-
stratum remaining, the sponge having completely destroyed it. Leidy (1890) states
that New Jersey fishermen call the y-stage colonies “bay pumpkins.” Long Island
Sound oystermen refer to the free-living colonies as ‘‘porpoise dung.”

In life, the y-stage colonies are golden yellow in color (Maerz and Paul, 1950,
Pl. 10, H-5, K-5, L-6) externally, the tubules being somewhat darker when con-
tracted. The endosome varies from deep chrome (op. cit., Pl. 9, L-7) to hazel (op.
Cit FA. 13 9-8).

SYSTEMATIC STUDIES Wi,

The shell perforations through which the incurrent papillae protrude vary
from 0.8 to 2.5 mm. in diameter; those through which oscular papillae protrude
are 2.0 to 4.5 mm. in diameter.

The general morphology of Cliona celata has been well described by ‘Topsent
(1900), Vosmaer (1933; this work must be read critically, however, because of Vos-
maer’s broad concept of C. celata), and Volz (1939).

European authors (Topsent, 1900, and Vosmaer, 1933) have recorded the
presence of spirasters in young colonies of C. celata, and Hopkins (1956a) reports
these spicules in some specimens of this species from Louisiana. Microscleres have
not been found in New England specimens of C. celata regardless of age. Simi-
larly, the oxeas reported by Topsent and Vosmaer (loc. cit.) in occasional colonies
of C. celata have not been found in American specimens, including the Gulf
Coast populations studied by Hopkins. Table 5 compares spicule sizes of C. celata
from Long Island Sound with those reported from other localities on the Ameri-
can Atlantic Coast.

Eggs observed in Long Island Sound specimens of Cliona celata are spherical,
varying from 32 to 38u in diameter. The nucleus varies from 14 to 16y in diame-
ter; the nucleolus is 5u. Topsent (1911) reports that the eggs of European speci-
mens measure 45.

DISTRIBUTION IN NortH AMERICA: Gulf of St. Lawrence to South Caro-
lina; Gulf Coast of Louisiana and Texas; Pacific Coast of North America.

Gulf of St. Lawrence (Lambe, 1900b, p. 164); Prince Edward Island (Lambe,
1896, p. 202; Old, 1941, p. 12); Mount Desert Island, Maine (Procter, 1933, p.
114); Casco Bay, Maine (Kingsley, 1901, p. 161, as Cliona sulphurea); Portland
Harbor, Maine (Verrill im: Verrill and Smith, 1873, p. 744); Vineyard Sound,
Mass., 11-18.5 meters (Desor, 1851, p. 68, as Spongia sulphurea); Vineyard Sound,
Mass., 2 to 30 meters (Verrill, 1871, p. 359, as Spongia sulphurea Desor; Verrill
in: Verrill and Smith, 1873, p. 744 and 1880, p. 232, as Cliona sulphurea); Vine-
yard Sound and Buzzards Bay, Mass., 4 to 35 meters (Sumner et al., 1913, p. 557);
Woods Hole, Mass. (Allee, 1923, p. 175; de Laubenfels, 1949, p. 23); Block Island
Sound, R. I., 20 to 40 meters (Hartman); off Long Island (Verrill, 1878, p. 406, as
Cliona sulphurea); Long Island Sound, low water to 28 meters (Verrill in: Verrill
and Smith, 1873, p. 744; Old, 1941, p. 12; Hartman); Great Egg Harbor, N. J.
(Verrill in: Verrill and Smith, 1873, p. 744, as Cliona sulphurea); Great Egg Har-
bor and Little Egg Harbor, N. J. (Leidy, 1857, pp. 162-163; 1890, p. 70; ‘Topsent,
1887, p. 9); Delaware Bay (Old, 1941, p. 12); Ocean City and Sinnepuxent Bay,
Maryland (Old, 1941, p. 12); Chincoteague Bay, Virginia (Old, 1941, p. 13);
Chesapeake Bay (Old, 1941, pp. 12-13); Ft. Macon, N. C. (Coues and Yarrow,
1879, p. 312); Beaufort Harbor, N. C. (George and Wilson, 1919, p. 138; Mc-
Dougall, 1943, p. 331; de Laubenfels, 1947, p. 34); South Carolina (Lunz, 1935,
p- 2; Hopkins, 1956b, p. 20); St. George’s Sound, Florida (Menzel, 1956, p. 1);
west coast of Florida (Carter, 1885, p. 207: = C. caribboea?); Louisiana (Moore,
1899) p: 93; Gary, 1906a;, p. 28; 190Gb, p. 50; 19074, p28; 1907b, p. 52; all as
Cliona sulphurea; Hopkins 1956a, p. 49); Aransas Bay, Texas, 114 meters (Hart-
man; also unverified records by Hopkins, 1956a, p. 54); California (de Lauben-
fels, 1932b, p. 47; Hartman, in: Smith et al., 1954, p. 19).

TABLETS
SPICULE DIMENSIONS OF CLIONA CELATA

1932b

LOCALITY GROWTH RANGE OR RANGE AND MEAN OF
AUTHOR STAGE LENGTH X WIDTH*—TYLOSTYLES
Prince Edward Island Lambe, 1896 a 229-320u x 4.9u
Mt. Desert Island, Me. | Procter, 1933 a 243-324-364
(mode)
Heads usually subterminal
Woods Hole, Mass. de Laubenfels,
1949 a 300u x 10u

Long Island Sound Old, 1941 a 220-400u x 4-10u
and Chesapeake By

Off Momauguin, Hartman a 193-286-369
East Haven, Conn. x 4.1-7.1-10.3yu
YPM #767

New Haven Harbor, Hartman a 189-274-357
Connecticut x 4.1-6.2-8.2u
YPM #835

New Haven Harbor, Hartman a 172-288-332
Connecticut x 5.3-7.6-10. 7p
YPM 4813

Off Stratford Pt., Hartman a 176-307-414
Connecticut x 4.1-7.2-9 .4y
YPM #766

Off Momauguin, Hartman B 213-296-385
East Haven, Conn. x 4.1-8.4-11.1p
YPM #767

Off Momauguin, Hartman y 213-323-377
East Haven, Conn. x 7.0-9.1-11.9u
YPM #767

Beaufort Harbor, George and a 200—400u x 4-9u
North Carolina Wilson, 1919

Beaufort County, Hartman a 238-350-426u
South Carolina x 8.2-12.5-17.2p
YPM #1859

Parris Island, Hartman a 221-306-398
South Carolina x 7.0-9.5-12.3y
YPM #1860

California de Laubenfels, a 200-270 x 2-9u

* Measurements reported by Hartman are based on 50 spicules per specimen.

18

SYSTEMATIC STUDIES 19
Cliona lobata Hancock, 1849

SynonyMy: See Topsent (1900, pp. 70-71) and Vosmaer (1953, pp. 362-364).

Discussion: Cliona lobata is another common boring sponge on the oyster
beds of Long Island Sound. The shell perforations produced by C. lobata are
slightly larger (incurrent, 0.2 to 0.5 mm.; excurrent, 0.8 to 1.6 mm.) than those
made by C. vastifica, but these two species are easily confused in the field. Cliona
lobata sometimes produces perforations in a reticulate pattern. The color of the
colonies is pale yellow.

The morphology of this sponge has been described by Topsent (1900) and
Volz (1939). The spicules (fig. 4) are (1) tylostyles which may have subterminal or
multiple heads and (2) spirasters of two categories—(a) those with spines arranged
in a spiral pattern around the spicule, the spines being more prominent at the
angles and ends of the spicule and (b) those with more abundant spines distrib-
uted at random over the surface. Spicules of the latter category are generally
smaller in size than the former. Topsent (1900) has figured the two types of spir-
asters.

Vosmaer (1933) pointed out the similarity of colonies of Cliona lobata to
young spiraster-containing colonies of Cliona celata found in European waters,

A O 20p

Ficure 4. Spicules of Cliona lobata. A. Tylostyles. B. Spirasters with spines in
spiral pattern. C. Spirasters with spines more evenly distributed. New Haven Har-
bor, Conn. YPM #1852L.

and inquired if “we are simply dealing with small, enfeebled specimens of C.
celata.” He relegated lobata to synonymy with celata. Topsent (1900), on the
other hand, had emphasized two differences between the two species: (1) C. lobata

20

MARINE SPONGES

TABLE 6

SPICULE DIMENSIONS OF CLIONA LOBATA

RANGE OR RANGE AND MEAN OF LENGTH X WIDTH

SPIRASTERS*
LOCALITY Spines
TYLOSIVLES arranged Spines evenly
in spiral distributed
pattern
New Haven Harbor, 156-198-234 26-43 7.9-17 .1-24.5y
Connecticut x 2.5-3.5 -4. 1 x 2.6-5.3u | x 1.8-2.7-3.9u
YPM #813 (8%) (92%)
New Haven Harbor, 172-197-226u 26-39u 7.0-17 .1-26.3yu
Connecticut x 2.1-3.1-4.1yu x. 3.08 x 1.8-2.3-3.5u
YPM #1852A (6%) (94%)
New Haven Harbor, 168—208-250u 26-50u 14.0-18 .0-21.Ou
Connecticut x 1.6-2.8-4. 1p x 3.7-4.4u | x 1.8-2.5-3.7y
YPM #1852C (16%) (84%)
New Haven Harbor, 152-191-217y 37-39u 10.5-18 .6—24.5y
Connecticut x 1.2-2.4-3.7y x 3.9-5.3u | x 1.6-2.7-3.9u
YPM #1852D (6%) (94%)
New Haven Harbor, 168-195-230 30-35 10.0-17 .0-21.0u
Connecticut x 2.1-3.4-4.1y x 3.5-4.4u | x 0.9-2.5-3.9u
YPM #1852E (6%) (94%)
New Haven Harbor, 148-180-209 28-44 14.0-19 .0-26.0u
Connecticut x 1.2-2.0-2.9u x 3.5-5.3u | x 1.4-2.5-3.5y
YPM #1852L (12%) (88%)
Chesapeake Bay 150—250u x 3-4 15-50pu x 2yu

(Old, 1941)

* All measurements (except those of Old) based on 50 spicules of each category per specimen.

has smaller papillae protruding from smaller perforations, and (2) it has a high
frequency of trilobed tylostyles. Vosmaer (1933) discounted the importance of
the latter character because such variant types of tylostyles are also found in other
clionids. In American waters, Cliona lobata is readily distinguishable from C.
celata, however. The papillae and shell perforations of the former show no over-
lap in size with those of the latter; C. Jobata always has an abundance of spirasters,
while C. celata always lacks these spicules, in New England waters at least.

DISTRIBUTION IN NorRTH AMERICA: Long Island Sound to South Carolina;
Louisiana; ‘Texas (?); California.

New Haven Harbor and vicinity, Conn., 20 meters (Hartman); Cold Spring
Harbor, Great South Bay, Long Island, N. Y. (Old, 1941, p. 13); Chesapeake Bay

SYSTEMATIC STUDIES 21

(Old, 1941, p. 13); Chincoteague Bay, Virginia (Old, 1941, p. 13); Beaufort Har-
bor, N. C. (Old, 1941, p. 13); Beaufort Co., S. C. (Hartman, specimen received
from Mr. Nathan Bowman); South Carolina (Hopkins, 1956b, p. 20); Louisiana
(Hopkins, 1956a, p. 49); Texas (unverified record by Hopkins, 1956a, p. 54);
California (Hartman, in: Smith et al., 1954, p. 19).

Cliona vastifica Hancock, 1849

Synonymy: See Lendenfeld (1898, p. 86, under the name Vioa vastifica); Top-
sent (1900, pp. 56-57); and Vosmaer (1933, pp. 402-411).

Discussion: This species of boring sponge is found commonly on oyster beds
in Long Island Sound, occurring from mean low water to depths of at least 10
meters. It is less common than C. celata and never, to my knowledge, overgrows
its substratum completely to form free-living colonies as is true of celata.

The morphology of Cliona vastifica has been described by Nasonov (1883, as
Cliona stationis); Lendenfeld (1898); Topsent (1900); Vosmaer (1933); and Volz
(1939). The spicules (fig. 5) are tylostyles which sometimes have subterminal or
multiple heads; acanthoxeas, which are often more abundant than the tylostyles;
and spirasters. Ihe oxeas are usually spined but occasionally smooth or spined

FicurE 5. Spicules of Cliona vastifica. A. Tylostyles. B. Smooth and microspined
oxeas. C. Smooth and microspined spirasters. (To left of C is a microspined style.)
New Haven Harbor, Conn. YPM #834.

29 MARINE SPONGES

only at the ends. In Long Island Sound specimens, the oxeas generally have a
slight central swelling; a few lack this characteristic. In specimens from the Bay
of Fundy examined by the writer a large percentage of the oxeas likewise show
traces of a central swelling; on the other hand, in specimens from South Caro-
lina (received from Mr. Nathan Bowman) only about 30 per cent of the oxeas
exhibit central swellings. Topsent (1900) figures several acanthoxeas with marked
central swellings from French specimens of vastifica, but other European authors
have not reported them.

New England colonies of C. vastifica are light yellow in color when alive. The
shell perforations through which the incurrent papillae protrude vary from 0.2
to 0.4 mm. in diameter; those through which the excurrent papillae protrude
vary from 0.6 to 1.4 mm. in diameter. The shell perforations of colonies living
in thin shells (such as Pecten magellanicus) are often arranged in reticulate pat-
terns; this is seldom the case in oyster shells, where the perforations are spaced at
random.

Table 7 compares spicule measurements of Long Island Sound specimens with
those from other localities on the North American Coast.

DISTRIBUTION IN NorTH AMERICA: From the Bay of Fundy to South Caro-
lina; Gulf of Mexico—northwest coast of Florida, coasts of Louisiana and
Texas (?), and Gulf of Campeche, Mexico.

Off Grand Manan Island, N.B. (Hartman); Fipennies Ledge, Gulf of Maine,
70 meters (Hartman); Stellwagen Bank, Massachusetts Bay, 40 meters (Hart-
man); Block Island Sound, 20-40 meters (Hartman); Long Island Sound, mean
low water to 10 meters (Hartman; Old, 1941, p. 13); York River, Chesapeake Bay
(Old, 1941, p. 13); Beaufort Harbor, N. C. (Old, 1941, p. 13); South Carolina
(Lunz, 1935, p. 2; Hopkins, 1956b, p. 20); Apalachicola Bay, Florida (Pearse and
Wharton, 1938, p. 635); Louisiana (Hopkins, 1956a, p. 49); Texas (unverified re-
port by Hopkins, 1956a, p. 54); Campeche Bank, Mexico (Topsent, 1889, p. 35).

Cliona truitti Old, 1941

Discussion: This species has not been found by the present writer along the
Connecticut shores of Long Island Sound, although Old (1941) reports it from
the north shore of Long Island, New York. It is a brackish water form differen-
tiated from vastifica in regard to the sizes and shapes of oxeas and spirasters. The
oxeas are somewhat longer than those of vastifica and almost always have a cen-
tral swelling which is frequently pronounced and may be multiple. As has been
pointed out previously, however, traces of a central swelling are characteristic of
most of the oxeas of C. vastifica in specimens from the New Haven region and
the Bay of Fundy. ‘Topsent (1900) has figured oxeas from French specimens of
vastifica in which the central swellings are as well developed as those in C. truitti.
The microspines on the oxeas of truztti are smaller, and the percentage of smooth
oxeas is higher than in vastifica. The spirasters of C. truzttz are considerably
smaller and less contorted than those of vastifica.

It is possible that the brackish water populations which Old has named
truittt are simply ecophenotypes of vastifica, a thesis supported by the occurrence
of similar variants in brackish waters in other parts of the world (see Part III).
On the other hand, Old’s distribution data give no indication of gradual varia-
tion with decreasing salinity; the boundary between the ranges of the two species
is very sharp, especially in Chesapeake Bay (fig. 44), with no zone of overlap. An

SYSTEMATIC STUDIES 93

TABLE 7
SPICULE DIMENSIONS OF CLIONA VASTIFICA AND CLIONA TRUITTI

RANGE OR RANGE AND MEAN OF LENGTH X WIDTH

LOCALITY AUTHOR
Tylostyles* Oxeas** Spirasters*
Chiona vastifica:

Grand Manan Isl., | Hartman | 164-223-295 62-93-131u 9 .0-13.7-20.3u
N.B. x 3.7-4.9-7.4u | x 2.1-3.8-6.2u | x 1.7-2.43.8y
YPM #893

New Haven Harbor,| Hartman | 139-202-250yu 70-114-164u 9.0-13.1-18.8u
Conn. x 2.9-4.0-5.7u | x 2.9-5.2-8.6u | x 0.9-2.2-3.0u
YPM #834

New Haven Harbor,| Hartman | 144-201-234u 57-108-148u 7.5-12.1-16.5y
Conn. x 2.1-3.44.5u | x 2.1-4.3-6.2u | x 1.5-2.2-3.8u
YPM #1851A

New Haven Harbor,| Hartman | 176—220-279y 57-100-156u 9.0-13.0-16.5u
Conn. x 2.5-3.8-4.9u | x 2.14.2-7.8u | x 1.7-2.3-3.0u
YPM #1851B

Chesapeake Bay Old, 1941 | 160—250yu 50-110u 6-23

x 3-5u x 2.5-4.0u x 1.0-3.5y
Georgetown Co., Hartman | 168-207-246 62-100-135yu 9.0-12.3-18.Ou
we: x 2.1-4.1-5.7 | x 2.1-3.8-5.3u | x 1.4-1.8-2.7p
YPM #1855

Beaufort Co., Hartman | 180—213-258u 82-117-160yu 9.0-12.2-17.3u
Se x 3.7-4.6-7.0u | x 2.9-4.1-5.3u | x 1.5-2.3-3.2p
YPM #1856

Cliona truttti:
Chesapeake Bay (?) | Old, 1941 | 190-225u 110-130 7-12u
x 2.5-3.5u x 4-5y x 0.5-2.0u

Charlestown Co., Hartman | 160-206-262 90-127-168yu 6.0-10.4-15.0pu
Suc) x 3.3-4.3-5.7u | x 2.9-4.0-5.3y | x 1.5-2.2-3.2y
YPM #1854

* Measurements reported by Hartman based on 50 spicules per specimen.
** Measurements reported by Hartman based on 100 spicules per specimen.

investigation of this matter through transplantation experiments would be of
interest. Hopkins (1956a, 1956b) reports an overlap in the ranges of Cliona vas-
tifica and C. truitti in Louisiana and South Carolina, however, virtually ruling
out this hypothesis. It is of interest to note, nevertheless, that in South Carolina
Hopkins (1956b) reports intermediates between Cliona vastifica and C. spirilla,
another brackish water species.

24 MARINE SPONGES

In the only specimen of this species available to me (from South Carolina),
the shell perforations made by the incurrent papillae vary from 0.1 to 0.3 mm.
in diameter; those made by the oscular papillae vary from 0.4 to 0.6 mm.

DIsTRIBUTION IN NorTH America: Long Island Sound to South Carolina;
Louisiana; Texas (?).

Long Island Sourd (Old, 1941, p. 13); Chesapeake Bay (Old, 1941, p. 13);
Choptank River, Patuxent River, Potomac River, St. Mary’s River, Wicomico
River, Sinnepuxent Bay, Maryland (Old, 1941, p. 13); Rantowles Creek, Charles-
ton County, S. C. (Hartman, specimen received from Mr. Nathan Bowman);
South Carolina (Hopkins, 1956b, p. 20); Louisiana (Hopkins, 1956a, p. 49); ‘Texas
(unverified record by Hopkins, 1956a, p. 54).

ORDER HALICHONDRINA
FAMILY HALICHONDRIIDAE Gray
Halichondria bowerbanki Burton, 1930b

SYNONYMY:
Halichondria coalita, Johnston, 1842, p. 135, Pl. XII, fig. 1.
Halichondria coalita, Bowerbank, 1866, p. 238; 1874, p. 102, Pl. XLI, figs.
18-20.
Halichondria bowerbanki Burton, 1930b, p. 489. See this paper for further
details.

SHAPE AND SIZE OF COLONIES: Young colonies form thin encrustations (Pl. 4,
figs. 4, 5) on rocks and algae. Colonies growing on the undersides of rocks retain
an encrusting form, the oscules not at all or only slightly raised above the gen-
eral surface of the colonies. Colonies growing in more exposed situations soon
develop vertical outgrowths which may be low tubules bearing terminal oscules
or lamellate ridges with oscula at their summits. Older colonies are variable in
shape but can be reduced to three general types: (1) colonies consisting of masses
of anastomosing branches, the branches sometimes being very thin (1-3 mm.; PI.
4, figs. 6, 7, 8) but usually thicker (up to 6-8 mm.; PI. 4, fig. 1). The largest colony
of this type in my collections measures 8 cm. high and 8 x 5 cm. across. (2) Col-
onies with a massive base from which arise vertical branches, more or less rounded
in cross section, irregularly placed (Pl. 3, figs. 7, 8 and PI. 4, fig. 3), sometimes
branching again or anastomosing with neighboring branches (PI. 3, figs. 4, 5). ‘The
largest colony of this type in my collections measures 7 cm. high and spreads out
16 cm. in one direction, 9 cm. in the other. (3) Colonies with a massive base from
which arise lamellate ridges, the distal edges of which grow out into flattened
processes (Pl. 3, figs. 6, 9 through 12). Largest colony collected: 6 cm. high and
9 x 9 cm. across. In some cases the lamellate processes grow up from the base of
the colony in a circular pattern to form “chimneys” (PI. 4, fig. 2) showing some
resemblance to colonies of H. panicea figured by Bowerbank (1874, p. 39, figs.
1, 2). In some colonies both flattened and rounded branches are found (PI. 3,
fies. 1092513).

Individuals studied at the Plymouth Laboratory, England, and the Station
Zoologique, Séte, France, fall into the second category listed above. Most of the
colonies collected in Europe have a massive base from which rounded, vertical
branches arise. One colony dredged off Plymouth, England, is made up of numer-
ous anastomosing branches.

CoLoR IN LIFE: Colonies of this species vary considerably in color. In sum-

SYSTEMATIC STUDIES 25

mer, the colonies are various shades of orange brown, cinnamon, topaz, yellow
beige, hazel, and bronze. (Maerz and Paul, 1950, Pl. 12, F-7, G-5, J-7, J-8, L-8;
Pl. 13, F-6, G-6, J-7, J-8, J-9; Pl. 14, I-9). Colonies containing zoochlorellae are
olive green (Maerz and Paul, 1950, Pl. 15, J-2). Winter specimens are darker in
hue, varying from cinnamon to drab, or in the case of specimens with zoochlorel-
lae, to olive drab and olive brown (Maerz and Paul, 1950, Pl. 12, F-7, H-7; Pl. 13,
D-4, F-4; Pl. 14, E-4, E-5, H-3; Pl. 15, H-5, J-7). Some colonies degenerate in part
during the winter, Josing their flagellated chambers, and becoming lighter in
color. Such colonies are apricot, buff, or cinnamon in color (Pl. 10, 1-7, J-7; Pl.
11, G-7, G-8; Pl. 12, F-9, G-7, H-7).

Specimens examined at Plymouth, England, tended to be dark beige to light
cinnamon in color (Maerz and Paul, 1950, Pl. 10, D-3, F-4, F-5; Pl. 11, D-4, E-5;
Pl. 12, D-6). The tips of the branches tended to be somewhat lighter (often near
Pl. 10, D-2).

ConsIsTENCy: Moderately firm and compressible.

SuRFACE: The surface of colonies of this species is irregular, being raised into
low mounds and ridges. In some specimens occasional spicule tracts pierce the
surface slightly, raising the dermis into low conules. This is usually not the case,
however.

OscuLEs: The oscules of Halichondria bowerbanki are highly variable in size
and position. They are sometimes borne on the summits of branches, but are also
distributed along the sides of branches and over the general surface of the colony.
Not all vertical branches bear terminal oscules. The oscules are elliptical or circu-
lar in outline, the dimensions of the axes varying from 135 x 100 to 5.0 x 2.5
mm. The majority vary from 1.0 to 2.5 mm. in diameter. Any one colony may
have oscules of widely different sizes. Although most of the oscular measurements
were made on preserved specimens, a similar range of sizes was observed in living
specimens examined.

Burton (1930b) states that the oscules of H. bowerbanki are inconspicuous,
perhaps because they were closed in the specimens examined by him. Although I
was unable to study Burton’s type specimen, all individuals which I collected at
Plymouth, England, and Séte, France, have the same range of oscular size and
position as those described above from the American populations.

PorEs: The pores, elliptical to circular in outline, show some tendency to-
ward forming vaguely defined groups on the surface. Pore size variation in three
specimens follows:

YPM #847. Pine Orchard, Conn. 33 x 33p to 20 x 20u (mean of 7, 29 x

25)

YPM +849. Hammonasset, Conn. 54 x 4Iu to 16 x l6u (mean of 50,
SI x21)

YPM +860. Double Beach, Conn. 68 x 43 to 13 x 9u (mean of 50,
36 x 21y)

EcrosOMAL ANATOMY: The dermal membrane is made up of a layer of thin
exopinacocytes. In thin sections perpendicular to the surface of the colonies, the
dermis is seen to lie above extensive subdermal lacunae belonging to both the
incurrent and excurrent systems of canals. Surface views of intact specimens re-
veal a system of such subdermal canals converging upon each oscule. By way of
these subdermal canals, water in the interior of the sponge finds its way to the
oscules.

26 MARINE SPONGES

The dermis is perforated by pores arranged in ill-defined groups and leading
into large subdermal incurrent lacunae. Trabeculae of amoebocytes reinforced
with spicules reach the surface at intervals and support the dermal membrane
above the extensive subdermal lacunae.

ENDOSOMAL ANATOMY: The interior of the sponge is complicated in structure.
Thin sheets or membranes of amoebocytes, anastomosing one with another,
separate the extensive interlocking systems of incurrent and excurrent canals. In
thin sections, the interior of the sponge has a very porous, open structure, with
extensive systems of canals separated by thin membranes of cells. ‘The flagellated
chambers are located in the cellular membranes and communicate between the
excurrent and incurrent canal systems. The flagellated chambers are ellipsoidal
in shape with dimensions varying from 30 x 25u to 45 x 35yu. The choanocytes
vary from 2.5 to 3.54 across. ‘The chambers appear to be eurypylous, but I am
not certain of this at present.

Although most specimens have the loose, open structure described above,
some have a denser endosome, the trabeculae of cells being more extensive. The
trabeculae are perforated by thin, incurrent canals which open into the flagellated
chambers, and these in turn lead into wide excurrent channels.

SKELETON: The chief components of the skeleton are tracts of oxeas running
toward the surface and anastomosing at intervals. ‘These tracts contain from sev-
eral to more than a dozen spicules in cross-section. They occur in the cellular
trabeculae which are also reinforced by many spicules of the same sort randomly
distributed. Spongin is scarce, occurring in small patches along the spicule tracts.

In many specimens spicule tracts are rare and ill-defined or entirely absent.
In these cases all the spicules are arranged at random in the cellular trabeculae.

The dermal spiculation repeats in essence the pattern of the endosomal skele-
ton. Multispicular tracts, running essentially parallel to one another, occasionally
anastomose to divide the dermis up into areas | to 2 mm. long and 200 to 300u
wide. In each dermal area bounded by multispicular tracts are imbedded numer-
ous single spicules which intermesh in such a way as to divide the area into many
smaller triangular or trapezoidal areas in which the pores open (fig. 8; Pl. 2, fig.
2). In specimens which lack endosomal multispicular tracts such tracts are also
weakly developed in the dermis, with intermeshing single spicules making up the
surface pattern in the main.

The spicules (fig. 6) are oxeas which taper gradually from the midpoint and
rapidly toward the extremities. For example, a spicule 280u in length was found
to measure 9u in thickness midway along its length; its thickness 90 to 100u in
either direction away from the midpoint was 8u; the last 40 to 50u at either end
of the spicule tapered rapidly to a point. Variant categories of spicules, such as
styles or strongyles, are only occasionally met with in this species.

The spicules are straight or gently curved at the midpoint; in some specimens
the central curvature is more pronounced. The spicules vary considerably in
size, the mean value for the length and width of the oxeas of 16 New England
specimens being (50 measurements per specimen): 285 x 7.6u. The range of the
means is: length, 231 to 328u; width, 5.7 to 9.8u. The over-all range of size is:
length, 135 to 390u; width, 3.7 to 13.1u.

The oxeas of European specimens studied are considerably larger than those
of American specimens. Spicule size ranges and means* of six European speci-
mens follow:

* Measurements based on 50 spicules per specimen.

SYSTEMATIC STUDIES 27

\
B
a)
50p
A
EE |
Sop

Ficure 6. Spicules of Halichondria bowerbanki. A. Oxeas and styles. New Haven,
Conn. YPM #814. B. Oxeas. Double Beach (Branford), Conn. YPM #1819.
Figure 7. Larva of Halichondria bowerbanki. Milford Harbor, Conn. (Stained
whole mount.)

Plymouth, England
YPM No. 2167A: length, 315-388-472u; width, 4-10-14y.
YPM No. 2167B: length, 265-390-515y; width, 6-11-15.
YPM No. 2168: length, 329-452-601; width, 8-/2-18y.
YPM No. 2169: length, 350-452-572u; width, 6-13-17y.
Range and mean values of all Plymouth specimens:
length, 265-421-601; width, 4-12-18.
Etang de Thau, Séte, France
YPM No. 2165: length, 307-380-458; width, 6-10-13y.
YPM No. 2166: length, 307-380-472; width, 3-6—7y.
Range and mean values of all specimens from Séte:
length, 307-380-472; width, 3-8-13u.

The spicules of the dermis and endosome show no conspicuous size difference
in New England specimens. Mean values for lengths and widths of oxeas of nine
Long Island Sound specimens are: dermis, 2891 x 7.74; endosome, 285u x
7.7u. Three specimens from southern Massachusetts reveal similar data: dermis,
261u x 6.34; endosome, 264u x 6.5. (See Table 9 for detailed data.) A specimen
from Séte, France, is consistent with the American populations in this character.
Mean spicule dimensions (length x width) follow: dermis, 3764 x 64; endosome,

28 MARINE SPONGES

TABLE 8
SPICULE DIMENSIONS OF HALICHONDRIA BOWERBANKI

SPICULE * SPICULE *
LENGTH* SPICULE WIDTH LENGTH* SPICULE WIDTH
LOCALITY RANGE AND RANGE SND. RANGE AND Cae
MEAN Lone MEAN ee
DERMIS DERMIS ENDOSOME ENDOSOME
Hammonasset, Conn. | 238-289-336u | 5.3-8.0-12.3u | 221-278-332u | 4.1-7.3-9.Ou
YPM #849
Double Beach 246-288-353 | 6.9-8.6-10.3u | 213-287-349u | 3.7-8.0-11.1p
(Branford), Conn.
YPM #1819
Double Beach 180-306-365 | 4.5-8.9-11.5u | 213-307-390u | 4.5-8.6-12.3y
(Branford), Conn.
YPM #1867
Double Beach 176-294-357u | 4.1-8.5-11.94 | 205-309-357u | 6.2-9.1-11.5y
(Branford), Conn.
YPM #1881D
Double Beach 246-293-349 | 4.1-6.2-9.8u 213-308-381 | 4.5-7.1-11.5y
(Branford), Conn.
YPM #763
Double Beach 197-246-316 | 4.1-5.7-7.8u 197-253-332p | 4.1-5.7-8.2y
(Branford), Conn.
YPM #840
Double Beach 180-276-336u | 4.5-8.0-11.9u 185-272-353 | 4.1-7.7-10.3y
(Branford), Conn.
YPM #860
Lighthouse Pt., 250-288-328 | 4.5-7.4-8.6u 193-296-336u | 4.1-7.3-8.6pu
New Haven, Conn.
YPM #814
Bass River, Mass. 217-260-303u | 4.5-7.0-8.2u 180—260-312u | 4.1-6.6-8. 6p
YPM #939
Lagoon Pond, 217-260-328u | 4.1-5.9-8.2u 209-272-353u | 4.1-6.5-8.2pu
Martha’s Vineyard,
Mass. YPM #940A
Lagoon Pond, 205-263-308u | 4.1-6.0-8.2u 193-259-295u | 4.5-6.#8.2u
Martha’s Vineyard,
Mass. YPM #940B
* All measurements based on 50 spicules per specimen. (Continued )

SYSTEMATIC STUDIES

TABLE 8—Conclusion

29

SPICULE DIMENSIONS OF HALICHONDRIA PANICEA

LOCALITY

Kent Isl., N. B.
YPM #919

Kent Isl., N. B.
YPM #903

Kent Isl., N. B.
YPM #867

Barter Isl., Maine
YPM #1243

Barter Isl., Maine
YPM #1244

Barter Isl., Maine
YPM #1246

St. Valéry-en-Caux,

France
YPM #1074

St. Valéry-en-Caux,

France
YPM #1075

Etretat, France
YPM #1077

Roscoff, France
YPM #2111A

Roscoff, France
YPM #2111B

Plymouth, England

YPM #2097A

Plymouth, England

YPM #2097B

SPICULE
LENGTH*
RANGE AND
MEAN
DERMIS

156-237-287

SPICULE WIDTH*
RANGE AND
MEAN
DERMIS

3.7-5 .0-7 .8u

SPICULE
LENGTH*
RANGE AND
MEAN
ENDOSOME

189-255-303

SPICULE WIDTH*
RANGE AND
MEAN
ENDOSOME

4.1-5.6-7.8pu

———EEaEa|ES|S]jSE]__ _ SE EOEOEeEeEEE—————_—

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180-224-303

4.1-6.2-9.0u

* All measurements based on 50 spicules per specimen.

164-262-328.

4.5-7.5-11.9p

30 MARINE SPONGES

383 x 6. On the other hand, in a specimen from Plymouth, England, the
dermal spicules are significantly larger than the endosomal ones. Mean dimensions
are: dermis, 4254 x 134; endosome, 389u x 11.

Specimens from Connecticut collected in winter and early spring tend to have
larger spicules than those collected in summer and early autumn. The ranges and
mean length and width measurements of oxeas of specimens collected in winter
and summer follow:

Winter (5 specimens): length, 176—300-390u width, 3.7-8.7-13.1.
Summer (11 specimens): length, 135-276-361y; width, 4.1—7.0-12.3p.

Eccs AND LARVAE: The eggs and larvae of H. bowerbanki appear in mid- and
late summer in New England. The eggs, subspherical in shape, measure 40 x 37y
to 45 x 40u in diameter; the nuclei are 16 x 12u; the nucleoli, 54 in diameter.
Larvae are released from late August (first seen leaving oscules of colonies in Con-
necticut on Aug. 21) through late November (see Part II). The larvae (fig. 7) are
obovoid in shape, 180 to 290u in length, 100 to 115u across the widest portion.
They are completely flagellated, with the longest flagella (31) at the anterior end.
At the posterior end is a zone of shorter flagella (81). In the interior of the stereo-
gastrulae, at the time of their release, are numerous small oxeas, with length and
width measurements varying as follows: 128 to 150u x 2.5 to 3.0u.

WINTER CONDITION: During the winter, when water temperatures are near
0° C., many colonies of this species remain alive and active in Connecticut; flagel-
lated chambers are present. Some, however, undergo partial degeneration, chang-
ing color as noted above, and losing their flagellated cells. Such colonies are com-
posed chiefly of masses of amoebocytes. In May, flagellated chambers reappear and
the colonies resume an active life. There is no evidence of gemmule formation in
this species.

DIsTRIBUTION IN AMERICA: Halichondria bowerbanki is an abundant sponge
along the Connecticut coast of Long Island Sound. It occurs from about half a
foot above mean low water down into subtidal waters. Occasional specimens can
be found in tide pools up to the mid-tide region. It is commonly found attached
to sea weeds and also grows on the upper surfaces, sides and undersides of rocks.
I have collected it intertidally at Hammonasset State Park (Meig’s Point); Stony
Creek; Pine Orchard; Double Beach (Branford); Lighthouse Point (New Haven);
Bradley Point (West Haven); and Milford. It also occurs in the offshore waters
of Long Island Sound down to depths of at least ten meters. It is not found in
the deeper waters of Block Island Sound.

Its range extends northward to the southern shores of Massachusetts. There
I have collected it in Lagoon Pond, Martha’s Vineyard, and at Bass River, Cape
Cod. Verrill (¢n: Verrill and Smith, 1873, p. 743) recorded it from Long Island
Sound near New Haven and from Vineyard Sound, Mass., as “Halichondria?
species undetermined, b.”

Discuss1on: Halichondria bowerbanki differs from H. panicea in the following
characters which will be discussed in greater detail below: 1) external form; 2)
dermal spiculation, both in arrangement and size of spicules; 3) larval structure
and season of reproduction. In other characters, such as spicule form and dimen-
sions, pore size, and oscular size, the two species overlap.

A comparative study has been made of specimens of H. bowerbanki from
southern New England; Plymouth, England; and Séte, France, and of specimens

31

SYSTEMATIC STUDIES

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SYSTEMATIC STUDIES 33

of H. panicea from Plymouth, England; Roscoff and St. Valéry-en-Caux, France;
and the coasts of Maine and New Brunswick in North America. In such a study,
one is immediately impressed by differences in external form. Mature colonies
of H. bowerbanki tend to be highly ramose, although some are flabellate. Young
colonies of this species (PI. 4, figs. 4, 5) are encrusting and some show a tendency
to produce many low oscular tubules; such colonies show some resemblance to
colonies of H. panicea figured by Bowerbank (1874, PI. 39, figs. 4, 5). But the type
of encrusting colonies of H. panicea with rows of oscules raised on ridges or
tubules, as figured by Bowerbank (1874, Pl. 40, figs. 3, 4) and represented in the
Peabody Museum collections by specimens from France are not found in popula-
tions of H. bowerbanki. Older encrusting colonies of H. panicea frequently bear
more or less regularly spaced taller oscular tubules (see, e.g., Johnston, 1842, Pl.
11, fig. 5; Bowerbank, 1874, Pl. 40, fig. 2; Vosmaer, 1933-35, PI. 59, fig. 10; Burton,
1928, fig. la; specimens in the Peabody Museum collections from Plymouth, Eng-
land, and the coast of Maine). Colonies of this form are lacking in H. bowerbanki,
in which the branches of older specimens are taller and much more erratic in
growth. The “‘cockscomb” variety of H. panicea figured by Bowerbank (1874, PI.
40, fig. 5) is approached by flabellate colonies of H. bowerbanki (PI. 3, fig. 11),
but these lack the apical rows of papillae shown in Bowerbank’s figure. One speci-
men of H. bowerbanki (P1. 4, fig. 2) has a chimney-like outgrowth resembling those
figured by Bowerbank (1874, PI. 39, fig. 1) and Burton (1928, fig. 1d) in H. panicea;
in H. bowerbank1, however, the chimney is not solitary but is located in the cen-
ter of a colony otherwise showing the branching habit characteristic of the species.
The young branching colonies of H. panicea illustrated by Renouf (1936, fig. 2)
do show a striking resemblance to colonies of H. bowerbanki. Regrettably Renouf
gives no structural details to indicate whether his specimens overlap in other
characteristics with populations of H. bowerbankt. The curious colony type with
long, thin branches figured by Vosmaer (1933-35, Pl. 60, fig. 4) and Grentzenberg
(1891, fig. 2) is represented by several specimens from Connecticut which, how-
ever, show a greater tendency toward anastomosis of the branches (PI. 4, figs.
6, 7, 8).

In summary, it may be said that in regard to colony form, the two species,
H. panicea and H. bowerbanki, overlap to some extent, but that some of the col-
ony shapes most characteristic of H. panicea are not represented in populations of
H. bowerbanki. The typical branching form of H. bowerbanki, on the other hand,
is not found regularly in H. panicea.

A striking difference between H. panicea and H. bowerbanki is seen in the
dermal skeleton. Surface views of H. panicea reveal a very regular network of
multispicular tracts which divide the dermis into chiefly oblong areas, in which
the closely spaced pores open (fig. 9; Pl. 2, fig. 1). Occasional isolated spicules lie
across the areas. The spicule tracts are from 30 to 90u in width and the dermal
areas between them vary from 90 to 150u in greater diameter. This very char-
acteristic pattern is apparent in specimens which I have examined from Plymouth,
England; Roscoff, France; and the coast of Maine.

The dermal spiculation of H. bowerbanki, as described above, is quite different
(fig. 8; Pl. 2, fig. 2). Multispicular tracts, when present, are widely spaced and
divide up the dermis into larger areas, 1 to 2 mm. long and 200 to 300u wide.
These areas are further subdivided by a pattern of overlapping individual
spicules. The pores are widely spaced. This is the most useful character for sep-
arating these two species; in my experience, it has proved consistent and reliable

34 MARINE SPONGES

for both American and European populations. Burton (1930b) comments that the
dermal reticulation of H. bowerbanki is more strongly marked than that of
H. panicea.

The dermal spicules of H. bowerbanki are approximately the same size as the
endosomal spicules. One specimen from Plymouth, England, is anomalous in this
regard, however, and has dermal spicules which are noticeably larger than those of
the endosome. In specimens of H. panicea from both New England and Europe,
the dermal spicules are consistently smaller than those of the endosome. Data for
these characters are summarized in Table 9. Fifty measurements were made for
each specimen. A detailed tabulation of these data is given in Table 8.

Vosmaer (1933-35) gives an excellent account of the ectosomal anatomy of
H. panicea, pointing out the variation which occurs. In all specimens cellular
pillars reinforced with tracts of spicules run to the dermis from the endosome.
These spicular tracts fan out at the surface to join the dermal tracts. The ecto-
somal skeletal architecture of H. panicea suggested to Dr. G. P. Bidder “the
vaulted interior of a Gothic crypt” (personal communication). In some specimens
the pores open into small subdermal cavities separated by pillars of cells and
these cavities open below into much larger lacunae or “crypts” (Vosmaer’s term)
leading into narrower inhalant canals which ramify the endosome. In other speci-
mens the small subdermal cavities are absent, and the pores lead directly into
large lacunae.

The ectosomal anatomy of H. bowerbanki resembles the latter arrangement,
with the pores opening directly into extensive lacunae which spread out horizon-
tally beneath the dermis. ‘The pillars of cells and spicules supporting the dermis
are thin and are generally reinforced by relatively few spicules. Bidder’s simile is
hardly appropriate for the weak skeletal architecture in the ectosome of H.
bowerbanki.

Other conspicuous differences between H. bowerbanki and H. panicea concern
the structure of the larvae and the season of reproduction. Data comparing the two
species in these regards are given in Table 10. See also figures 7 and 33.

Chronic lumpers such as Vosmaer (1933-35) would have little trouble in in-
cluding H. bowerbanki in a broad concept of H. panicea. ‘There is no doubt that
the two species overlap in many characteristics. Both species are highly variable in
structure and form and hence separating them is a difficult matter. The discus-
sion above focuses attention on a series of characters which, in my experience,
represent consistent differences between the species and on which I have based
my conclusion that the two species are separate and distinct. The dermal skeletal
pattern and the structure of the larvae seem to be the most reliable distinguishing
features. Vosmaer, however, was even reluctant to accept larval differences as being
significant. He states (1933-35), “I for my part have no doubt that the larvae will
be found just as variable as the adult sponges.”

Although the shape and appearance of colonies of H. bowerbanki from Amer-
ica and Europe show good agreement, as does the structure of the dermal skeleton,
specimens from populations on opposite sides of the Atlantic do exhibit certain
differences. It has already been pointed out that the spicules of specimens from
England and France are significantly larger than those of New England colonies.
There is also a difference in larval structure, bearing out in some measure the
comment of Vosmaer quoted above. The larvae of H. bowerbanki from Roscoff,
France, figured by Topsent (1911), are larger than those found by the present
writer in specimens from the Connecticut population. The pattern of the larval

SYSTEMATIC STUDIES

55

Ficure 8. Portion of dermis of Halichondria bowerbanki showing characteristic
arrangement of skeleton and pores. Lightly shaded areas represent openings from
subdermal cavities into channels leading to the interior of the sponge. More
heavily shaded areas represent trabeculae of cells extending from dermis to in-
terior. Lagoon Pond, Martha’s Vineyard, Mass. YPM +940.

~V3
SS

ama

Ficure 9. Portion of dermis of Halichondria panicea showing characteristic ar-
rangement of skeleton and pores. Plymouth, England. YPM +2097.

36 MARINE SPONGES

flagella also differs. The posterior flagella of Topsent’s specimens form a long
tuft, while those of Connecticut larvae are short. See Table 10 and figure 7. The
breeding seasons are similar on both sides of the Atlantic. It is highly probable
that the American and European populations represent valid subspecies; sub-
specific names are not proposed at the present writing, since data from a larger
series of specimens from Europe are needed.

Some colonies of H. bowerbanki bear a resemblance in external form, spicule
dimensions and oscular size to Swartschewsky’s (1905) figure of Halichondria grossa
Schmidt. The former author makes no reference to the characters used in my
diagnosis of bowerbanki, however, and it is difficult to draw a detailed comparison
of the species. Swartschewsky states that the oxeas are arranged in a disorderly
fashion, only rarely being grouped into more or less long, thick clusters, running
in all directions in the endosome. This describes the condition of the endosomal
skeleton of some specimens of H. bowerbanki fairly well. Whether or not bower-
banki and grossa are synonyms cannot be answered on the basis of our present,
incomplete knowledge of the latter species.

Swartschewsky’s figure of Halichondria luxurians (Lieberkiihn) also resem-
bles certain colony types of H. bowerbanki. However, H. luxurtans has styles, tylo-
styles, and strongyles as spicules.

The taxonomic status of H. bowerbanki (which has been referred to as H.
coalita in most of the literature on the species) has been clarified by Burton
(1930b). The name “coalita” was first used by Miiller (1776, p. 256) who cited as
his type a specimen figured by Ellis (1755, p. 80, Pl. 32, fig. F). Burton has pointed
out correctly that Ellis’ figure represents a specimen of Haliclona oculata (Pallas,
1766). Burton has shown further that most subsequent references to coalita con-
cern Haliclona oculata or Halichondria panicea or are unrecognizable. ‘To my
knowledge Fleming (1828, p. 522) was the first author to refer coalita to the genus
Halichondria, but he cites Miiller’s (1776) original incorrect designation. The
name “coalita’” is thus unavailable for use with Halichondria. Johnston (1842,
p. 135) very probably had reference to the Halichondria in question, but he also
applied Miiller’s name. Bowerbank (1866, 1874) likewise used the name, “coalita,”
citing Johnston as the author of the species. Bowerbank’s specimen was a gift
from Grant whose many references (1825, 1826a, 1826c, 1827) to Spongia coalita
without indication doubtless concern the species under discussion.

Bowerbank (1866, p. 239) suggests that Esper’s (1794, Pl. 41) Spongia suberosa
“is undoubtedly the same species as that designated Spongia coalita by Dr. Grant,
and Halichondria coalita by Johnston.” I have been unable to consult Esper’s
paper at this writing, but the possibility exists that swberosa is an available name
for the species in question. However, Ehlers (1870) does not mention this species
in his restudy of Esper’s sponges, and it is probable that the specimen is lost.
Indeed, Bowerbank’s specimen of Halichondria coalita is apparently the only one
still extant of those described by early authors. Burton has wisely chosen this
specimen, figured by Bowerbank in 1874 (PI. 41, fig. 18), as the type for his new
species, Halichondria bowerbanki.

ORDER POECILOSCLERINA
FamILy MICROCIONIDAE Hentschel
Microciona prolifera (Ellis and Solander, 1786) Verrill, 1873
SYNONYMY:
Spongia prolifera Ellis and Solander, 1786, p. 189

SYSTEMATIC STUDIES 37

Spong|ija ostracina Rafinesque, 1819, p. 150
[non] Spongia prolifera Grant, 1826a, pp. 115, 116, 123, and 1827, pp. 135,
138

Spongia urceolata Desor, 1851, p. 67
Microciona prolifera, Verrill, in: Verrill and Smith, 1873, p. 741
Clathria delicata Lambe, 1896, p. 192

Discussion: This well known sponge is abundant in Long Island Sound where
it grows commonly on oysters, beginning life as a thin encrustation. As the sponge
ages, vertical finger-like lobes develop (PI. 4, fig. 11); in old colonies these increase
in number and anastomose to form bushlike colonies with the branches intricately
interwoven (PI. 4, fig. 9). Such colonies often reach a large size; the largest one
collected on an oyster bed by the present writer measures 20 cm. in height and
spreads out 25 cm. in one direction and 8 cm. in the other. Often flattened
branches arise from the basal encrustation, producing fan-shaped structures which
break up into lobes distally (Pl. 4, fig. 10, 13). In extreme cases, such flattened
branches form cup-shaped colonies by growing up from the substratum in a cir-
cular pattern (PI. 4, fig. 14). Desor (1851) also mentions this colony type.

M. prolifera occurs from mean low water down to depths of at least ten meters
in the New Haven area. It is occasionally found in tide pools up to two feet
above mean low water. Intertidally this sponge grows on the upper surfaces, the
sides, and the undersides of rocks. Intertidal individuals are chiefly encrusting or
have lobes no more than two or three cm. in height. The sponge is perennial in
growth in Long Island Sound in both offshore and intertidal locations, but indi-
viduals growing in the latter environment undergo considerable degeneration
during the winter and die back to thin encrustations which renew growth in the
spring. Large, bushy colonies which require several years of growth to assume
this form, seldom, if ever, are found near mean low water.

The color of M. prolifera varies seasonally and with exposure to light. In the
summer and autumn most specimens are tomato red (Maerz and Paul, 1950, PI.
3, H-12) or terra cotta (Pl. 4, D-12, E-12, and H-12) in color; others are burnt
orange (PI. 3, F-12) or burnt sienna (PI. 5, G-12). In winter and spring many speci-
mens are browner in color, near gold brown (PI. 14, E-10, B-11; Pl. 15, E-10) or
henna (PI. 6, G-12); others approach burnt sienna (PI. 5, J-12) or terra cotta (PI. 4,
B-12). The redder hues are characteristic of colonies exposed to the light; colonies
growing on the undersides of rocks are nearer orange brown in color.

The general morphology of this sponge has been described by Wilson (1912)
and George and Wilson (1919). The present writer has little to add, except to
point out the occurrence of cords of elongate cells in which the developing
spicules are aligned. These structures were noted in small regenerating colonies
fixed and stained by Patricia J. Harris, of the Bingham Oceanographic Labora-
tory. Cell cords of this general nature are apparently of wide occurrence in the
Demospongiae, having been noted by Herlant-Meewis (1949) in Spongilla and
Suberites domunculus and by the present author in several species of Haliclona
as well as Microciona.

Table 11 compares the sizes of spicules from Long Island Sound specimens
with those reported from other localities along the American Atlantic Coast.
Three categories of megascleres (fig. 10A) occur: (1) The most abundant ones are
long, stout, subtylostyles (less frequently styles) with smooth shafts and spiny,
or occasionally smooth, heads (the heads may be unspined). ‘These spicules project

MARINE SPONGES

38

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50p

FicureE 10. Spicules of Microciona prolifera. A. Megascleres (see text). B. Toxas.
C. Isochelas. New Haven, Conn. YPM #754. Isochelas from specimen from Double
Beach (Branford), Conn. YPM #1912E.

laterally from the fibers of the sponge, forming tufts at the surface where the
fibers end. (2) ‘Thin subtylostyles and styles, usually with microspined heads.
Wilson (1912) and George and Wilson (1919) regard this category of spicules as
developmental stages. The shorter ones of these are distributed throughout the
sponge and doubtlessly do represent immature spicules. ‘The longer ones, how-
ever, are localized chiefly in tufts at the surface and probably represent a distinct
category. This hypothesis is supported by the gap in width measurements between
the thin and stout types of megascleres. (3) Short acanthosubtylostyles and
acanthostyles, in which both shafts and heads are spined, the latter more thickly.
These spicules stand out at right angles from the fibers (pointed ends outward)
in the interior of the sponge colony. In addition there are two categories of micro-
scleres: (1) palmate isochelas (fig. 10C) and (2) toxas (fig. 10B). The latter are
always common in specimens from the New Haven area, although they are rare
or absent in specimens north of Cape Cod. Most of the toxas in New Haven
colonies are very short (12-20u) but a few range up to 50-60u in length.

The available data suggest that the sizes of the short, spiny subtylostyles and
toxas increase in northern waters, but there is no apparent correlation of sizes
of the other megasclere types or chelas with latitude. George and Wilson (1919)
report an increase in width of the large subtylostyles in old portions of colonies.

SYSTEMATIC STUDIES 4]

Vosmaer (1933) regards Microciona prolifera as a cosmopolitan species, but
his extensive synonymies are highly questionable.

DIsTRIBUTION: Nova Scotia and Prince Edward Island to South Carolina,
Louisiana and Texas (?).

Prince Edward Island (Lambe, 1896, p. 12, as Clathria delicata); Nova Scotia
(Lambe, 1900b, p. 160, as Clathria delicata); Mt. Desert Isl., Me., 24m. (Procter,
1933, p. 104); Portland, Me. (Lambe, 1896, p. 192, as Clathria delicata); Bass
Rocks, Cape Ann, Mass., mean low water (Hartman); Edgartown Harbor,
Martha’s Vineyard, Mass., 8 meters (Desor, 1851, p. 67, as Spongia urceo-
lata); Vineyard Sound, Mass., low water to 20 meters (Verrill, in: Verrill and
Smith, 1873, p. 742; 1880, p. 232); Woods Hole, Mass., and vicinity, low water
toro) meters (Sumner ef al.) 1903, 2/5593 ‘Allee, “1923; p. 175;' Procter, 1933,
p- 104; de Laubenfels, 1949, p. 12); Long Island Sound, low water to 20 meters
(Verrill, in: Verrill and Smith, 1873, p. 742; 1880, p. 232; Hartman); shores
of Long Island (Rafinesque, 1819, p. 150, as Spongia ostracina); Great Egg
Harbor, N. J. (Verrill, in: Verrill and Smith, 1873, p. 742); coast of New Jer-
sey (Ellis and Solander, 1786, p. 189, as Spongia prolifera); Chesapeake Bay,
13 meters (Cowles, 1931, p. 329); Ft. Macon, N. C. (Verrill, in: Verrill and Smith,
1873, and Coues and Yarrow, 1879, p. 312); Beaufort Harbor, N. C. (Wilson,
1912, p. 3; George and Wilson, 1919, p. 157; McDougall, 1943, pp. 331-332; de
Laubenfels, 1947, p. 35); South Carolina (Verrill, in: Verrill and Smith, 1873,
p. 742; Hopkins, 1956b, p. 24); Louisiana and Texas (Hopkins, 1956a, p. 44,
records an encrusting Microciona from these states, but does not identify it as
prolifera.).

FAMILY MyxILLipaAE Hentschel
Lissodendoryx isodictyalis (Carter, 1882) Topsent, 1897

SYNONYMY:
Halichondria isodictyalis Carter, 1882, p. 285
Tedania leptoderma Topsent, 1889, p. 49
Lissodendoryx leptoderma, Topsent, 1894, p. 35
Lissodendoryx isodictyalis, Topsent, 1897, p. 456
Lissodendoryx carolinensis Wilson, 1912, p. 11

Discussion: This species is widely distributed in subtropical and tropical seas
(West Indies, Mediterranean Sea, and East Indies) and extends northward along
the Atlantic Coast of North America from the Florida Keys to Woods Hole, Mass.
It occurs chiefly as a fouling organism on wharf piles in the region from Beaufort,
North Carolina, to Woods Hole, and has been collected near mean low water
as an encrustation on rocks at Double Beach (Branford), Connecticut. .

The morphology of this sponge has been described by Wilson (1912, p. 11),
George and Wilson (1919, p. 150), and de Laubenfels (1936, p. 93). Variations in
spicule dimensions are presented in Table 12. Specimens (carolinensis type) from
the Atlantic Coast of North America from Beaufort, N. C. to Woods Hole are
characterized by the presence of large sigmas (mean chord length, 29.3u) and
small chelas (mean chord length, 18.9) in contrast to specimens (isodictyalis type)
from subtropical American waters which have small sigmas (mean chord length,
18.0u) and large chelas (mean chord length, 21.54). Both of these variations occur
together in the Mediterranean, however, and this observation is of questionable
value for differentiating two subspecies on the American coast.

42 MARINE SPONGES

One specimen of Lissodendoryx isodictyalis from Long Island Sound was
collected by Verrill in 1874. No more exact locality was given by Verrill, but it
is probable that the specimen came from the eastern end of the Sound, since
reference to the list of dredging stations of the U. S. Fish Commission for 1874
indicates that no stations were worked west of Saybrook, Connecticut, during
that year. Verrill’s specimen is massive in form and measures 42 x 30 x 15 mm.
in linear dimensions.

Recently L. isodictyalis has been found by the present writer at Double
Beach (Branford), Connecticut, growing as thin encrustations on rocks. Portions
of two colonies were collected in November, 1955; one was growing near the mean
low water level, another, about half a foot below. By January, 1956, the remainder
of these colonies had the form of elongate encrustations about 8 x 3 cm. in dimen-
sions. Their color in life was light olive tan (Maerz and Paul, 1950, Pl. 12, F-2,
G-2). In each, the surface was raised into numerous low knob-like projections
which in one case were no higher than 0.5 mm., in the other, about 2 mm. The
oscules were about 0.5 mm. in diameter. Spicule dimensions for these specimens
are given in Table 12. (See also fig. 11.) In August, 1956, another encrusting
colony was collected, 4.5 x 4.5 cm. in dimensions (see Pl. 4, fig. 12); this colony
is provided with numerous translucent papillae, 1.5 to 2.5 mm. high and 0.5 mm.
in diameter. A large colony, 8 x 8 cm., was observed at the same locality in Novem-
ber, 1956.

FicurE 11. Spicules of Lissodendoryx isodictyalis. A. Megascleres: styles and
tylotes. B. Sigmas. C. Isochelas. Long Island Sound. YPM #2117.

SYSTEMATIC STUDIES

TABLE 12
SPICULE DIMENSIONS OF LISSODENDORYX ISODICTYALIS

43

LOCALITY AND AUTHOR

Woods Hole, Mass.
(Hartman; specimen

from M. D. Burkenroad)

YPM #931**

Eastern Long Isl. Sound
(Hartman; A. E. Verrill

colls.)
VPM 2117

Double Beach, Conn.

(Hartman)
YPM #2037F-2*

Double Beach, Conn.

(Hartman)
YPM #2037F-1*

Beaufort, N. C.
(Hartman; specimen
from I. M. Newell)
YPM #616**

Beaufort, N. C.
(George and
Wilson, 1919)

Fort Jefferson, Fla.

(de Laubenfels, 1936)

Harrington Sound,
Bermuda
(Hartman; specimen
from W. Bergmann)
YPM #699**

Bermuda
(de Laubenfels, 1950)

RANGE OR RANGE AND MEAN OF
SPICULE LENGTH X WIDTH

Tylotes

RANGE OR RANGE AND

Sigmas

MEAN OF SPICULE LENGTH

Chelas

fn | |

136-147-163
x 3.7-4.4-6.6u

160-175-187u
x 3.3-4.0-4.8u

| | fa S

167-180-191
x 1.5-5.1-7.0n

Me OF). Le

ey

148-176-197
x 6.2-7.5-8.2u

164-183-197
x 4.1-5.0-6. 6p

—— | —_ —— — —— |_| |

139-163-185
x 6.1-7.6-8.2u

148-174-205
x 4.5-5.2-7.4u

162-243-316u
x 4.4-7.0-9.2u

159-178-191
x 4.4-5.5-7.3y

—————— — ————_ | — — — — —— § | |

x 5-8u

ROL

S$ | $s |§ |]

136-152-163
x 2.6-3.3-3.71

176-192-207
x 2.6-3.1-3. 7

| | |

150u x 3-5y

180u x 2-4

12-17-20

* Fifty of each category of megascleres measured; 25 of each category of microscleres measured.

** Twenty of each category of megascleres measured ;

measured.

10 of each category of microscleres

(Continued)

44 MARINE SPONGES

TABLE 12—Concluded
SPICULE DIMENSIONS OF LISSODENDORYX ISODICTYALIS

RANGE OR RANGE AND MEAN OF RANGE OR RANGE AND
SPICULE LENGTH x WIDTH MEAN OF SPICULE LENGTH
LOCALITY AND AUTHOR TEES Se cs
Styles Tylotes Sigmas Chelas
Acapulco, Mexico 168u x 6.3y 200u x 6.3y 17u 25
(Tyre: Carter, 1882)
Guadeloupe 160-175 x 3-6 | 198-2104 x 4-Sy | 25-30u 25-30
(Topsent, 1889)
Monaco 150-170yu 180-190u 28-32 14-16
(Topsent, 1925a)
Naples 230-280u 265-310u 25-31 29-33
(Topsent, 1925a) x 2-6u RO OL
Venice, Italy 215 200u 33-35, 55 | 25-27p
(Topsent, 1925a)
Amboina, Moluccas 180—200u 200-215 25-30 30-31
(Topsent, 1925a) x 5-7u x 5-6u
Celebes 200u x 5—6u 220u x 5—6u 22u 30u

(Topsent, 1925a)

DISTRIBUTION IN NORTH AMERICA AND THE CARIBBEAN SEA: Woods Hole,
Mass., to the Florida Keys; Bermuda; West Indies; Venezuela; Acapulco,
Mexico.

Woods Hole, Mass. (Hartman—specimen collected by M. D. Burkenroad);
eastern Long Island Sound (Hartman—specimen collected by A. E. Verrill);
Double Beach (Branford), Conn. (Hartman); Beaufort, N. C. (Wilson, 1912 pau
as L. carolinensis; George and Wilson, 1919, p. 150, as L. carolinensis; McDougall,
1943, p. 331, as L. carolinensis; de Laubenfels, 1947, p. 35; Hartman—specimens
collected by I. M. Newell); South Carolina (Hopkins, 1956b, p. 24); Fort Jeffer-
son, Florida (de Laubenfels, 1936, p. 93); Bermuda (de Laubenfels, 1950, p. 73);
Guadeloupe (Topsent, 1889, p. 49, as Tedania leptoderma); Puerto Cabello,
Venezuela (Carter, 1882, p. 285, as Halichondria isodictyalis); Acapulco, Mexico
(Carter, 1882, p. 285, as Halichondria isodictyalis).

ORDER HAPLOSCLERINA

FAMILY DESMACIDONIDAE Gray
Tsodictya Bowerbank 1863, emend.

The genus Jsodictya was first mentioned by Bowerbank in 1859 in a general
paper on the anatomy and physiology of sponges, in which he figured spicules

SYSTEMATIC STUDIES 45

of two species of the genus. A spicule of Isodictya anomala Bowerbank, MS
(stc) was figured to illustrate the “inflato-fusiformi-acerate” category of spicules
(pp. 286, 323, Pl. 23, fig. 11); a spicule of Isodictya lobata Bowerbank, MS (sic)
was figured to illustrate the “dentato-palmate inequi-anchorate” category of spic-
ules (p. 327, Pl. 24, fig. 58). No definition of either the genus Isodictya or
the species anomala and lobata was given by Bowerbank at that time. Thus, the
generic name, Jsodictya, was not validated in this early paper, although the
species anomala and lobata must stand from this date in accordance with Article
27a in the International Rules of Zoological Nomenclature.

Bowerbank defined and described Jsodictya in 1863 stating at this time that
“Tsodictya infundibuliformis is perhaps the most perfect type of the genus.” In
1864 he repeated the same definition of the genus, including the statement about
I. infundibuliformis and in addition, he clearly designated I. palmata and
normani as the type species of the genus. It follows that Bowerbank did not
intend to designate infundibuliformis as the type species by his reference to it
in 1863 but was using the word “type” in a different sense. Isodictya palmata,
therefore, is the type species of the genus by designation of the original author,
a conclusion accepted by Dendy (1924, p. 334). Burton (1936, p. 143) has implied
that Isodictya must be reserved for anomala and another name chosen for pal-
mata; this seems inadmissable to me.

Bowerbank included a large number of species in the genus Isodictya; most
of these have since been transferred to other genera. If the above comments are
accepted, the name /sodictya must be restricted to species allied to palmata. Lund-
beck’s definition of Homoeodictya (1905) is an acceptable one for Isodictya in
this sense, except for its restriction to forms with diactinal megascleres. Evidence
is presented in the following pages which argues for an extension of the con-
cept of Isodictya to include species with monactinal megascleres, otherwise an-
swering Lundbeck’s diagnosis of Homoeodictya. Such species have previously
comprised part of the genus Esperiopsis.

Isodictya deichmannae?® (de Laubenfels) new comb.
SYNONYMY:

Esperiopsis quatsinoensis Lambe, 1893. (As used by Procter, 1933, p. 94)
Neosperiopsis deichmanni de Laubenfels, 1949, p. 15

Discussion: De Laubenfels (1949) established a new genus and species for
this sponge which occurs along the Atlantic Coast of North America from Block
Island Sound to the Grand Banks off Newfoundland. For a generic character
he pointed to the presence of “distinctive” microscleres which he considered as
reduced chelas resembling “‘sigmas with forked ends.’’ His figures of these micro-
scleres are very crude and bear only a vague resemblance to the spicules observed
by the present author in de Laubenfels’ type specimen (MCZ 446910). The micro-
scleres (fig. 12b) found in all fifteen specimens examined by the writer cor-

‘respond to the description and excellent figures given by Lundbeck (1905) of
the “Homoeodictya” type of chela. This chela has the axis extended outward so
as to form a projection inside the tooth of each end. In side view immature forms
of this microsclere might possibly be interpreted as being sigmas with forked

9 De Laubenfels named this species in honor of Dr. Elisabeth Deichmann, but never-
theless used the masculine ending for the specific name, “deichmannt.” This lapsus calami
is corrected above in accordance with article 14c of the International Rules of Zoological
Nomenclature.

46 MARINE SPONGES

ends; however, specimens viewed from the front give unmistakable evidence of
the chelate form, as do completely formed specimens in either view. The micro-
scleres characteristic of this species are unquestionably identical with those of
the species Isodictya palmata (fig. 13b).

Indeed, this sponge bears a general resemblance to Isodictya palmata (La-
marck)!° Bowerbank, so much so that it is frequently impossible to distinguish
the two species on the basis of external characteristics alone (see PI. 6, figs. 1 and
4). The only certain difference between them is the type of megasclere present.
In one form, styles are present; in the other, oxeas. The similarities between the
two species are as follows:

(1) In both species a reticulate skeletal framework exists consisting of pri-
mary spicule tracts held together by a spongin sheath and radiating out
from the axis of the branches to the surface. Secondary fibers connect
the primary tracts at irregular intervals. (See figures 14 and 15.)

(2) In both species the same variety of isochelate microscleres exist.

(3) The oscules in both species are distributed along the sides of the
branches and are of similar diameters.

(4) The range of external shapes is similar in both species. (See Pl. 5, figs.
3, 4, 5, and Pl. 6, figs. 1-5.)

In spite of these many resemblances the two species have been placed in dif-
ferent genera by previous authors on the basis of the single criterion, megasclere
shape. Table 11 indicates that the two species overlap in regard to this character.
Specimens of Isodictya palmata from the Atlantic Coast of North America fre-
quently possess a small number of styles in addition to oxeas1! (fig. 13); specimens
of the styliferous species often bear some oxeas (fig. 12). There seems to be little
question but that these two species must be included in the same genus.

The question now arises as to whether the two forms belong to the same
species. On the basis of geographical distribution this is unlikely. [sodictya pal-
mata is a widely distributed boreal species which is known from northern Eng-
land, Scotland, the Shetland Islands, the Orkneys (these records from Johnston,
1842, and Bowerbank, 1866); and from the Sea of Okhotsk (Burton, 1935). Along
the Atlantic Coast of North America, it occurs off Nova Scotia and in the Gulf
of Maine south to Eastport, Maine (Lambe, 1896; Hartman). The style-bearing
form is restricted to the coast of North America from the Grand Banks to Block
Island Sound, according to present records based on specimens in the Peabody
Museum.

A related group of species occurs on the Pacific Coast of North America. These
were described by Lambe (1893) as belonging to four distinct species of the genus
Esperiopsis: rigida, vancouverensis, quatsinoensis, and laxa. From the descriptions,
figures, and ranges given in Lambe’s article it seems not unlikely that these four
species represent growth forms of a single species. A reéxamination of Lambe’s
types will be necessary to clarify this point. Procter (1933) who identified the style-
bearing Isodictya from Mt. Desert Island, Maine, as Esperiopsis quatsinoensis
Lambe, has tabulated the skeletal dimensions of Lambe’s E. vancouverensis, quat-

10'Topsent (1933) regards Lamarck’s Spongia palmata as equivalent to Isodictya
palmata.

USwartschewsky (1906) reports the presence of occasional styles in specimens of
I. palmata from the White Sea.

SYSTEMATIC STUDIES 47

—

Figure 12. Spicules of Isodictya deichmannae. A Megascleres: styles, subtylostyle,

strongyle, centrotylote oxea. B. Microscleres: isochelas in front (left and bottom)
and side (right) views. Kent Isl., N. B. YPM #890.

20p

FicurE 13. Spicules of Isodictya palmata. A Megascleres: oxeas, style, strongyle.
B. Microscleres: isochelas in front and side views. Kent Isl., N. B. YPM #923.

48 MARINE SPONGES

1a

<A

\ Si
A —
Xs
1
ane

|

, Vell
Na

—,

—-

LEE
SS—

=

——
—

Ficure 14. Portion of skeletal framework of Isodictya deichmannae. (Section per-
pendicular to surface.) Kent Isl., N. B. YPM +890.

sinoensis, and laxa. These three species intergrade in respect to these characters,
and Procter concluded that they are synonymous. E£. rigida differs in bearing two
categories of styles in respect to width. This same variation occurs in some speci-
mens of the East Coast species; these may represent specimens which were pre-
served at a time when active spicule-formation was taking place, the thin styles
probably representing growth stages.

Procter’s suggestion that the East and West Coast forms are identical is invalid
in the present author’s opinion. De Laubenfels’ separation of the eastern and
western species on the basis of megasclere width seems unjustified unless other
characters can be found to differentiate them. More conspicuous characteristics
separating the two species are the position and sizes of the oscules. All four of
Lambe’s species of Esperiopsis bear terminal oscules. Even E. quatsinoensis, which
is described and figured with rows of oscules down the sides of the branches, also
bears larger oscules at the tips of the branches. Two specimens of this species
from St. Lawrence Island, Alaska (AMNH Nos. 264 and 267) were examined by
the author. The colonies consist of clusters of branches each of which bears a
large terminal oscule. No oscules are present along the side walls of the tubular
branches. ‘The oscules are elliptical or circular in outline and vary from 12 mm. x
7 mm. to 3mm. x 3 mm. in diameter. On the other hand, all of the New England
specimens studied by the present author lack terminal oscules and instead have
the oscules arranged irregularly or in rows along the sides of the branches. The
oscules in these specimens are circular and are much smaller, varying in diameter
from 1 mm. to 3 mm.

De Laubenfels’ species, “deichmannae,” is therefore a valid one, though not on
the basis of the character suggested by him. The species is assigned to the genus

SYSTEMATIC STUDIES 49

=

fle

— |
—S==

<a

SS
—~s=

Ficure 15. Portion of skeletal framework of Isodictya palmata. (Section perpend1-
cular to surface.) Kent Isl., N. B. YPM 4923.

Isodictya, being virtually indistinguishable from I. palmata except in regard to
megasclere form.

It is of interest to note that microsclere form and size are remarkably constant
in the three species studied here (see Table 13). On the other hand, the megascleres
vary in length and width considerably; even their form is not constant in any one
individual. A certain percentage of diactinal megascleres regularly occur along
with the styles; small numbers of styles occur with oxeas. Hentschel (1929) has
pointed out a number of trends in megasclere form correlated with temperature.
Exotylote spiculation is an arctic phenomenon. Spicule size characteristically in-
creases in colder waters. Oxeas often replace styles in waters with lower tempera-
tures. The species pair under consideration here offers support for the two last
mentioned trends.

Isodictya palmata appears to be a widespread subarctic species which has given
rise to more southern species on both the East and West Coasts of North America.
On the East Coast, at least, the northern and southern species overlap in part of
their ranges.

DIsTRIBUTION OF I[sodictya palmata IN NorTH AMERICA: Nova Scotia to
Eastport, Maine.

Nova Scotia and Sable Island (Lambe, 1896, p. 190); Minas Basin, Bay of

50

SPICULE DIMENSIONS OF AMERICAN DESMACIDONIDS

LOCALITY

CATALOGUE NUMBER

Grand Banks
YPM #419**

Kent Island, N. B.
YPM #890*

Eastport, Maine
YPM #152**

Casco Bay, Maine
YPM #446**

Casco Bay, Maine
YPM #2122*

Georges Bank
YPM #2123**

Off Cape Cod
YPM #2124**

Nantucket Shoals
YPM #2125*

Off Nantucket
YPM #2112*

Off Nantucket
YPM #2126*

Off Nantucket
YPM #2127*

Off Nantucket
YPM #2113*

Woods Hole, Mass.
YPM #2128*

MARINE SPONGES
TABLE 13

MEGASCLERES

RANGE AND MEAN

CHELAST
RANGE AND

LENGTH AND WIDTH |MEAN LENGTH

Isodictya deichmannae

STYLES

131-196-231
x 9.5-11.7-14.3y

119-180-209
x 5.3-8.5-11.9p

146-191-239u
> a Ea Acro Aa

131-182-231
x 6.2-11.0-15.4y

119-179-201
x 4.1-9.9-14.4y

169-234-254
x 5.5-11.0-13.9p

139-185-316u
x 3.7-7.7-11.0p

119-178-205
x 4.1-8.8-14.4y

144-180-205
x 4.1—10.3-14.8y

103-148-185
x 2.5-6.4-13.5y

119-170-209
x 3.7-7.5-13.9u

98-160-197
x 3.3-10.2-15. 64

113-—160-179u
x 5.1-7.0-9.5u

OXEAS
25-28-33 —
18-23-31 —
21-24-33 Sao)
21-26-33 6:5
25-29-37 —
25-27-32, 1.0
22-26-29 0
19-21-30 2.0
23-26-30u —
14-19-23 4.0
Microscleres =
rare; range of
6: 12-18u
21-25-29u 4.0
23-26-29 —

* Megasclere measurements based on 100 spicules per specimen.
** Megasclere measurements based on 200 spicules per specimen.

t Measurements based on 25 spicules per specimen.

PERCENTAGE OF
OTHER MEGASCLERES

STRONGYLES

0.5

5.0

0.5

3.0

2.5

1.0

2.0

1.0

a

1.0

(Continued)

SYSTEMATIC STUDIES 51

TABLE 13—Concluded
SPICULE DIMENSIONS OF AMERICAN DESMACIDONIDS

MEGASCLERES CHELAST

LOCALITY RAE ni aieae PERCENTAGE OF
CATALOGUE NUMBER Sppbiegbieb 4 dis ANGE SAND OTHER MEGASCLERES

LENGTH AND WIDTH |MEAN LENGTH

Woods Hole, Mass. 123-160-185 20-26-30p. 3.0 —
MCZ #6910* x 3.7-6.7-8. 6p
Block Isl. Sd. 119-170-197 22-25—S0ofh 5.0 1.0

YPM #762* x 4.5-8§.1-12.3u

Block Isl. Sd. 108-159-216u 22-25-29u 120 —
YPM #817** x 7.0-8.4-10.2u

Noank, Conn. 139-182-216u 21-25-29u oo —

YPM #2129** x 5.5-8.8-12. 1p

Overall mean 177 x 8.8u 25 a -——

Tsodictya palmata

OXEAS CHELAS STYLES STRONGYLES

116-177-231p 23-28-33 0.5 aa
x 5.5—9.5-13 9p

Minas Basin, N. S.
YPM #2114**
169-229-277 24-26-33 1.0 —
x 8.4-11.3-14.6u

Kent Isl., N. B.
YPM #923**
162-219-262u 24-28-29 520 —
x 5.9-10.2-13.5y

Eastport, Maine
YPM #4**

172-216-258 23-26-29 me) —
x 4.5-8.3-16.0u

Eastport, Maine
YPM #4A*

Overall mean 210u x 9.8u 27 = ==

Esperiopsis quatsinoensts
STYLES CHELAS OXEAS STRONGYLES

St. Lawrence Isl., 172-192-220u 24-28-35 = aa

Alaska x 7.7-12.1-15.7p
AMNH #264f
St. Lawrence Isl., 154—-190-220u 22-25-29 — a
Alaska x 9.2-12.1-14.6yu
AMNH #267t

Overall mean 191y x 12.1 27u — aa

a

52 MARINE SPONGES

Fundy (Lambe, 1896, p. 190; Hartman—A. E. Verrill collections); off Kent Island,
N.B. (Hartman); off Eastport, Me. (Hartman—A. E. Verrill collections).

DIsTRIBUTION OF Isodictya deichmannae IN NortH America: Grand
Banks to Block Island Sound.

Grand Banks, exact locality unknown (Hartman—A. E. Verrill collections);
off Kent Island, N.B. (Hartman); Eastport, Me. (Hartman—A. E. Verrill collec-
tions); Mount Desert Region, Me. (Procter, 1933, p. 94, as Esperiopsis quat-
stnoensis); Casco Bay (Hartman—A. E. Verrill collections); Georges Bank (Hart-
man—A. E. Verrill collections); Stellwagen Bank, Massachusetts Bay, 34 meters
(Hartman); Vineyard Sound and Nantucket (Verrill im: Verrill and Smith, 1873,
p-. 742, as Isodictya sp.); Woods Hole, Mass. (de Laubenfels, 1949, p. 15, as Neo-
speriopsis deichmanni); Watch Hill, R. I. (Verrill in: Verrill and Smith, 1873,
p. 742, as Isodictya sp.); Block Island Sound, 20-40 meters (Hartman); off Noank,
Conn. (Hartman—A. E. Verrill collections).

It is impossible to say to which of the two species Sumner et al. (1913, p. 559)
were referring as “Desmacidon palmata (Johnston)” since they mention no micro-
scopic characters.

FAMILY HALICLONIDAE de Laubenfels
Haliclona oculata (Pallas, 1766) Grant,!? 1841

SyNonyMy: See Lundbeck, 1902, p. 9 and Arndt, 1935, p. 100. To be included
in the synonymy is Chalina arbuscula Verrill (in: Verrill and Smith, 1873, p. 742).

Discussion: Haliclona oculata (Pallas) is a common sponge along the Atlantic
Coast of North America from the Gulf of St. Lawrence to North Carolina. Verrill
(in: Verrill and Smith, 1873) pointed out that specimens of this species occurring
south of Cape Cod are “much more delicate, with more slender and rounder
branches” than northern specimens.!* He placed the southern forms in a new
species, Haliclona (his Chalina) arbuscula. Since this species, although well de-
scribed by Verrill (op. cit.) has given rise to some confusion in the literature, the
present writer has undertaken to clarify its status. A biometric analysis has been
made of specimens from a series of localities along the Atlantic Coast of North
America from the Bay of Fundy to Long Island Sound. Mean dimensions of four
characters (total height of colonies, mean width of branches at their bases [eight
measurements per colony] and mean spicule length and width [50 measurements
per colony] ) are summarized in Tables 14, 15, 16. On the basis of these data, it
is apparent that two morphologically distinct populations exist in the region
under consideration, one occurring north of Cape Cod, the other, south of the
Cape. The mean dimensions of each of the four characters for all localities con-
sidered collectively (1) to the north of and (2) to the south of Cape Cod, have
been determined and are listed in Table 17. The means of three of the characters
(total height, branch width, spicule length) differ significantly (P<0.01 in each
case) when the northern and southern populations are compared, although the
observed ranges overlap in all instances. The mean values of spicule width for
the two populations are not significantly different, although the largest values for

12 Grant spells the specific name as “occulata,’ but it is clear from his figure that he
had this species in mind.

13 Compare specimens from south of Cape Cod (PI. 7, figs. 1, 2; Pl. 8, figs. 1, 2, 3; Pl. 9,
fig. 5) with those occurring north of the Cape (PI. 9, fig. 2; Pl. 10, figs. 1, 2, 3).

SYSTEMATIC STUDIES 53

TABLE 14
VARIATIONS IN THE FORM OF HALICLONA OCULATA (PALLAS)

NO. OF RANGE AND MEAN RANGE AND MEAN BRANCH
LOCALITY SPECIMENS | HEIGHT OF COLONIES- WIDTH-MILLIMETERS*
MEASURED MILLIMETERS
Long Island Sound 14 100-156-190 (4+ 31) | 3.1 x 2.1-4.4x 2.8-
7.1x3.7 (+ 1.7, + 0.8)
Block Island Sound 11 170-267-355 (+ 59) | 4.3x 3.6-5.4x 3.6-
7.1x3.7 (4 1.6, + 0.7)
Vineyard Sound 5 155-199-260 (+ 34) | 2.9x 2.7-4.4x 3.1-
6.5x 3.0 (+ 1.8, + 0.8)
Massachusetts Bay 10 170-267-330 (+ 48) | 7.7x5.3-9.3 x 5.5
(Stellwagen Bank) 11.8x 6.9 (+ 2.7, + 1.4)
Gulf of Maine 9 170-277-420 (+89) | 5.2x 3.87.5 x 4.8
9.9x 4.6 (+ 2.5, + 1.1)
Georges Bank 1 290 9.0x 3.5-9.1 x 6.1-
1325 x 7.0) (41220), s- £ 25)
Atlantic Coast of N. A., 30 100-207-355 (+ 44) | 2.9x 2.7-4.7 x 3.2-
South of Cape Cod 7 m3: 7 (42157), 2057)
Atlantic Coast of N. A., 20 170-272-420 (+ 72) | 5.2x3.8-8.4x5.1-
North of Cape Cod 13.5x 7.0 (+ 2.6, + 1.3)
West Coast of Portugal 6 350 2.0-6.0
(Arndt, 1941)
Channel Coast of France 2 170-190-210 3.6x3.14.2x3.5-4.8x3.8
(St. Valéry-en-Caux) (+ 1.1, + 0.8)
Hastings, Sussex, England 1 ca. 170 ca. 4.0-5.0
(Bowerbank, 1874)
Faroes (Lundbeck, 1902) 2 s0572295t 1°) We ae Oa,

ee ee

* Eight measurements per colony.

this character were observed in specimens from north of Cape Cod. If the four
character means for colonies from each of the localities north of Cape Cod are
compared with one another, there is no significant difference among them; the
same is true for the localities south of Cape Cod (with the exception of total
colony height in which there is a significant difference between the Long Island
Sound and Block Island Sound populations).

Coefficients of difference (C. D.) for the four characters have been calculated,
comparing values for all colonies north and south of Cape Cod (Mayr, Linsley,
and Usinger, 1953); see Table 17. These data bear out Verrill’s observations that
the chief difference between the northern and southern populations is in branch
size. In view of the extent of overlap in other characters, however, it seems un-
warranted to separate the two populations into subspecies.

54 MARINE SPONGES

CABLES

SUMMARY OF VARIATIONS IN THE SPICULE SIZES OF HALICLONA
OCULATA (PALLAS)

NO. OF MEAN SPICULE LENGTH | MEAN SPICULE WIDTH
LOCALITY SPECIMENS| AND RANGE-MICRONS AND RANGE-MICRONS
MEASURED
Long Island Sound 6 84—116-156p (410.5) | 3.7-8.5-12.8u4 (41.6)
Block Island Sound 4 84-116-1394 (+ 8.0) | 2.9-7.7-11.0u (41.3)
Newport, Rhode Island 1 88-118-132u (4 7.9) | 5.5-9.2-11. 7p (41.4)
Vineyard Sound 3 77-107-137p (+ 9.6) | 2.9-6.2-11.7p (41.2)
Atlantic Coast of N. A., 14 77-114-156p (+ 9.6) | 2.9-7.8-12.8y (41.5)
South of Cape Cod
Massachusetts Bay 3 110-137-168 (10.9) | 4.8-9.7-12.8y (41.3)
(Stellwagen Bank)
Casco Bay 5 108—129-156u (+ 8.1) | 4.1-7.8-13.2u (41.3)
Bay of Fundy 2 103—-128-165p (413.4) | 7.3-10.1-12.8u (41.3)
Georges Bank 1 119-141-156p (+ 9.6) | 5.7-8.9-11. 5m (E1.1)
Nantucket Shoals 1 127—-145-168u (410.0) | 5.7-8.6-11.5u (40.9)
Atlantic Coast of N. A., 1) 103-133-168 (410.9) | 4.1-8.8-13.2u (41.3)
North of Cape Cod
West Coast of Portugal 6 83-103 u 8.0-10.0u
(Arndt, 1941)
Channel Coast of France 1 68—97-115y 3.7— 4.8- 5.5u
(St. Valéry-en-Caux)
Hastings, Sussex, England 1 ca. 119y ca. 9.4
(Bowerbank, 1874)
Faroes (Lundbeck, 1902) 2 120-149-178 8.0-11.0-13.0u

TABLE 16

SPICULE DIMENSIONS OF HALICLONA OCULATA*

LOCALITY

SPICULE LENGTH

SPICULE WIDTH

RANGE AND MEAN | RANGE AND MEAN

4.4-9.5-11.0u

New Haven, Conn. YPM #430 114-123-146p
Mansfield Pt. (East Haven), Conn. YPM #810 84-102-121y 4.8-7.3-10.6u
Thimble Islands, Conn. YPM #469 88-107-124u 5.9-7 .0-8. 1p

* All measurements based on 50 spicules per specimen.

(Continued)

SYSTEMATIC STUDIES 55

TABLE 16—Concluded
SPICULE DIMENSIONS OF HALICLONA OCULATA*

LOCALITY SPICULE LENGTH SPICULE WIDTH

Hammonasset, Conn. YPM #820 992-120-156 3.7-9.5-12.8u
Hammonasset, Conn. YPM #823 88—120-135u 3.7-9 .2-11.3y
Noank, Conn. YPM #2130 90-125-152y, 4.8-8.4-11.7y
Block Island Sound YPM #756 101-116-134 =
Block Island Sound YPM #757 95-107-117u 3.3-7 3-9 .9u
Block Island Sound YPM #773 84-116-135y, 2.9-7 .0-10.2u
Block Island Sound YPM #780 112-124-139 7.3-8 .8-11.0u
Off Newport, R. I. YPM #2131 88-118-132u 525 922-11 ip
Woods Hole, Mass. MCZ #6908 79-102-115u 2.9-5.1-6.6u
Woods Hole, Mass. YPM #942 77-103-119 3.7-5.5-8. 1p,
Vineyard Sound YPM #2132 97-116-137u 5.1-8.1-11.7p
Off Nantucket YPM #2137 127-145-168 5.7-8 .6-11.5u
Stellwagen Bank, Massachusetts Bay YPM #965A 110-731-156 5.5-9.2-12 .4y
Stellwagen Bank, Massachusetts Bay YPM #965B 110-134-163 4.8-9 .2-12.1p
Stellwagen Bank, Massachusetts Bay YPM #965C 115-145-168 7.3-10 .6-12 .8u
Casco Bay, Maine YPM #448 108—135-150p 5.9-9 .5-13.2u
Portland, Maine YPM #2116 115-130-144 5.3-9.1-12.3y
Portland, Maine YPM #2133 111-124-156 4.9-7 .§8-12.3y
Portland, Maine YPM #2135 111-126-152u 4.9-7 .5-10.7u
Portland, Maine YPM #2136 115-130-144 4.1-5.1-8.6pu
Georges Bank YPM #2138 119-141-156 5.7-8.9-11.5u
Kent Isl., N. B. YPM #924 110-136-165 7.3-10.2-11.Op
Kent Isl., N. B. YPM #901 103-120-137 7.3-9.9-12.8y

* All measurements based on 50 spicules per specimen.

RANGE AND MEAN | RANGE AND MEAN

56 MARINE SPONGES

TABLE, 17

COEFFICIENTS OF DIFFERENCE FOR MEASUREMENTS OF HALICLONA
OCULATA NORTH AND SOUTH OF CAPE COD

ee

MEANS AND S. D.
CHARACTER $$ C.D. PERCENT OF
OVERLAP

North of Cape Cod | South of Cape Cod

eee eee ee Oe — — — ——————————— | anc

Colony height 272mm. +72 156mm. + 31 1.126 87
Branch size* 44mm. +13.0 15mm. +5.7 1.545 94
Spicule length 133u +10.9 114u + 9.6 0.927 82
Spicule width 8.8u 41.3 7.8u+ 1.5 0.361 a

a —

* Product of the cross-sectional diameters.

The size differences between the northern and southern populations are clearly
correlated with water temperature, and represent but another example of a com-
mon phenomenon, that of increase in size in northern, colder waters. Hentschel
(1929) has pointed out a correlation of spicule length with water temperatures in
several genera of siliceous sponges. His observations are given in Table 18.

Maximum and minimum surface and bottom temperatures for the regions un-
der discussion are given in Table 19. The boundary between the northern and
southern populations coincides with the summer cold-water barrier in the region
of Cape Cod pointed out by Parr (1933). It is of interest to note that Vineyard
Sound specimens agree well in size with those from farther south, whereas the
one specimen available from Nantucket Shoals is close to the northern forms.
Bigelow (1933) has pointed out the existence of a zone of upwelling in the latter
area, which keeps the summer temperatures of Nantucket Shoals considerably
colder than those just to the west.

Verrill (in: Verrill and Smith, 1873) recorded an overlap in range between
Haliclona (Chalina) oculata and his Haliclona arbuscula, listing records of the

TABLE 18

SPICULE SIZES OF SILICEOUS SPONGES IN ARCTIC AND TROPICAL
REGIONS

(From Hentschel, 1929)

GENUS POLYMASTIA| MYCALE MYXILLA | |\GEEERGS
SPICULE TYPES* STYLES STYLES | ACANTHOSTYLES OXEAS
Arctic 1303 431 287 313
Tropical Regions of Atlantic 1140 365 157 264
Antarctic 1427 587 579 520

I

* Mean lengths of megascleres in microns.

SYSTEMATIC STUDIES 57

PABLE*19

TEMPERATURE RANGES AT LOCALITIES WHERE HALICLONA
OCULATA OCCURS

SURFACE] SURFACE

TEMPER-| TEMPER- BOTTOM BOTTOM
LOCALITY ATURE* | ATURE* | TEMPERATURE* | TEMPERATURE* AUTHOR
MAXI- | MINI- MAXIMUM MINIMUM
MUM MUM
Long Island Sound 20.5-21} 3.5 16-18.5 1-3.5 Fuglister, 1947
(30 meters) (30 meters) Riley, 1948
Block Island Sound 20.5-21} 3.5 15.5-19 3-5 Fuglister, 1947
(20-30 meters)| (20-30 meters)| Riley, 1948
Vineyard Sound 20-20.5} 3.5 15.5-20 1-3 Fuglister, 1947
(15-30 meters)| (15-30 meters)| Riley, 1948
Massachusetts Bay 18-> 20) 2-2.5 |} 11-12 1-2 Bigelow, 1928
(Stellwagen Bank) (20-25 meters)} (20-25 meters)
Gulf of Maine 14.5-15} 1-1.5 7-9 eS Bigelow, 1914,
(Casco Bay) (30 meters) (30 meters) 1928
Georges Bank 15-18 | 3-4 6-10 2.5-3.5 Bigelow, 1928
(120 meters) | (120 meters)
Kent Island 10-11 1.5-2.0} 9-10 1.5-2.0 Bigelow, 1928
(40 meters) (40 meters)
Nantucket Shoals 12-16 | 2.5-4.5| 12-14 3-4 Bigelow, 1933
(30-40 meters)| (30-40 meters)
Channel Coast of 17-17.5] 7-8 — — Hutchins and
France Scharff, 1947

(St. Valéry-en-Caux)

—_—— qx | —_ —q—| |) qq qe ——_——_——_.

Faroes 11.5-12| 6-6.5 — — Hutchins and
Scharff, 1947

* All temperatures recorded in degrees centigrade.

former from off Watch Hill, R. I., and off Gay Head, Martha’s Vineyard. It is
apparent from Table 14 that Block Island Sound specimens (see also Pl. 7, figs.
1, 2; Pl. 8, figs. 1, 3) are significantly taller than those from Long Island Sound
(Pl. 8, fig. 2; Pl. 9, fig. 5) and in regard to this character resemble the northern
populations. But in branch size and spicule dimensions, the Block Island Sound
specimens agree with other populations south of Cape Cod. Verrill was probably
misled by the large colony size when he regarded the Watch Hill specimens as
being identical to forms from the Gulf of Maine.

The populations of H. oculata in Vineyard Sound and off the western end of
Martha’s Vineyard are distinctive in colony form and color. Verrill (op. cit.,

58 MARINE SPONGES

p. 497) describes specimens from off Gay Head as having flattened stalks which
fork distally and divide into numerous digitate branches.14 The color of these
individuals is described as dull orange red when alive. Dr. Werner Bergmann has
collected specimens identical with these in shallow water off Woods Hole, Massa-
chusetts. One specimen of his collection available to the author (Pl. 9, fig. 4)
shows the following spicule dimensions: length (range and mean), 76.9-103.2—
119.04; width (range and mean), 3.7—5.5-8.lu. The total height of this colony is
185 mm. and the mean branch width is 6.5 x 3.0 mm. This specimen thus fits into
the southern population as far as the dimensions of the four characters listed above
are concerned. It seems to differ from typical representatives of this population in
two ways; (1) its palmate manner of branching with terminal branches arising
from broad, flattened bases; (2) its orange-red color. The range of variation in
form in this species is great. Some specimens from Block Island Sound and Long
Island Sound show a similar tendency to a formation of palmate branches, and it
is probable that all intermediates could be found between the typical mode of
branching from the base of the sponge to the palmate condition. On the other
hand, the orange-red color has never been observed by the present writer in any
specimens of either the northern or southern populations. It is of interest that
this distinctive color, observed by both Verrill and Bergmann in much the same
region, should be associated in both cases with the extreme palmate branching
habit.1 It will be necessary to obtain a larger series of specimens from this region,
as well as cytological information and data on breeding habits and larval char-
acteristics, in order to determine the status of this population. For the time being
it is regarded by the present writer as a variant population of H. oculata, close to,
but not identical with, populations to the south.

It is of interest to note that, at least as far as the few observations and litera-
ture references available to the present author are concerned, a similar correla-
tion of structural characteristics with temperature is present in Haliclona oculata
in European waters. Spicule measurements of two specimens of Haliclona oculata
from the Faroes, as reported by Lundbeck (1902), agree with those of the northern
population of the western North Atlantic. On the other hand, specimens collected
for the writer by French fishermen at St. Valéry-en-Caux, on the northern coast
of France (PI. 9, fig. 3), show colony and spicule dimensions comparable to those
of the southern population of the North American Coast. (See Tables 14, 15.) A
specimen figured by Bowerbank (1874, Pl. 66) and collected in the English Chan-

14 Short colonies with a palmate pattern of branching (PI. 9, fig. 1) were collected by
Verrill off Nantucket. In some of these colonies the flattened basal portions break up into
many thin branches distally, thus resembling the colonies described by Verrill from off
Gay Head. More exact locality data for these colonies are not recorded, but they are
probably from the relatively shallow waters of Nantucket Sound rather than from the
Shoals to the south of the island. ‘They bear a close resemblance externally to the Woods
Hole and Vineyard Sound populations.

15 Sumner et al. (1913, p. 558) report that Dr. J. A. Cushman was uncertain of the
identity of the “Chalinas’” in the Woods Hole area and preferred to enter all of them as
“undetermined.” De Laubenfels (1949, p. 32) suggests that Verrill’s Vineyard Sound speci-
mens of Chalina arbuscula may represent Haliclona palmata (Ellis and Solander); how-
ever, the colony height and spicule dimensions given by de Laubenfels for a specimen of
H. palmata from Woods Hole are much smaller than those of Verrill’s Woods Hole speci-
mens of Chalina arbuscula (see Pl. 9, fig. 1A). Burton (1930a, p. 511) regards Spongia
palmata Ellis and Solander as “clearly an uncommon form of Chalina oculata occasionally
occurring on the British coasts.”

SYSTEMATIC STUDIES 59

nel off Hastings fits into the same group. The English Channel specimens, al-
though resembling the more southern American population in having small
spicule dimensions and branch width measurements, differ from the latter in a
tendency for the oscules to be raised above the surface of the branches on small
conules.

Specimens of Haliclona oculata from the west coast of Portugal have round
branches; the spicules are somewhat shorter than those of either the southern
population of the American Coast or the English Channel specimens. Arndt (1941)
regarded the Portuguese population as a subspecies, tavarest.

A word must be said about Fristedt’s records of “Chalina arbuscula Verrill”
from the southwest coast of Sweden (1885) and from northern Spitsbergen, north-
ern Siberia, and the Bering Straits (1887). These records, which have been cited
by Hentschel (1929) and Arndt (1935), if true, would invalidate the discussion
of the zoogeography of Haliclona oculata presented here. Alander (1942) has
concluded that the forms which Fristedt identified as “Chalina arbuscula Ver-
rill” actually belong to several species including Haliclona implexa (Schmidt)
and Haliclona montagui (Fleming). Apparently Fristedt’s specimens are not sy-
nonymous with the species studied by Verrill in the western North Atlantic.

It is impossible to say at present whether the size differences observed between
the northern and southern populations have a genetic or ecophenotypic basis.
Transplantation experiments have so far proved impossible because of the diffi-
culty of transplanting the northern specimens successfully to Long Island Sound.
Experiments on the effect of temperature on spicule size and spongin content
would be of interest. The sharp temperature break separating the two popula-
tions in the region of Cape Cod may isolate them reproductively, but definite
evidence of this is lacking. Specimens from Block Island Sound bearing larvae
were collected on July 23, 1944, but breeding colonies from north of Cape Cod
have not been found as yet. It the Block Island Sound colony, eggs measure
53 x 40u; embryos, from 230 x 230u to 245 x 185u.

Nores oN MorPHoLocy: Although Verrill’s description (in: Verrill and Smith,
1873, pp. 742-43) of the southern population is adequate and the northern popu-
lation is well-known from descriptions of northern European representatives of
the species, several comments on the morphology of Haliclona oculata are worthy
of mention. The amount of spongin associated with the skeleton varies greatly
from specimen to specimen and shows no clear-cut correlation with geographical
distribution. Figures 16, 17, and 18 illustrate the range in variation in the south-
ern population. Here the skeletal framework consists of a rectangular network
of single spicules and these are sometimes imbedded in continuous fibers of
spongin, sometimes joined together at their tips only by small amounts of spon-
gin. In the northern population (fig. 19) spongin fibers are the rule, with the
main tracts bearing from two to four spicules per cross-section, and the connect-
ing fibers usually containing single spicules. Mesh sizes vary with spicule length.
The amount of spongin joining the spicules may vary in different parts of the
same colony. Figure 20 illustrates different portions of a colony from the Channel
Coast of France. In this case, there is a basic network of spicule-containing spongin
fibers running through the branches, but between these fibers runs a secondary
reticulation of spicules joined at their ends by small amounts of spongin. Solid
fibers are more common in the interior of the branches.

In colonies from both the northern and southern American populations are
found a few extensive longitudinal spongin fibers, almost or entirely devoid of

60 MARINE SPONGES

Figure 16. Haliclona oculata. Portion of skeleton (section perpendicular to sur-
face) with small amount of spongin. Spongin stippled. Long Island Sound. YPM
#2118.

FicurE 17. Haliclona oculata. Portion of skeleton (section perpendicular to sur-
face) with spongin tracts. Spongin stippled. Block Island Sound, 35 meters. YPM
ge 101.

spicules. They are illustrated in a specimen from Long Island Sound (fig. 18).
The regular network of spicules and spongin found through the sponge colony
is joined to these occasional, sturdy spongin fibers which serve to give additional
support to the skeleton.

Width measurements of the fibers in question for four specimens from north
and south of Cape Cod are as follows: Long Island Sound (Off Mansfield Pt.,
Conn.), 25 to 50u; Long Island Sound, 23-35u; Casco Bay, 30-50u; Bay of Fundy
(off Kent Island, N.B.), 33-46u. Heavy fibers of this type are not present in speci-
mens from the Channel Coast of France.

Tracts of elongate cells in which spicules are aligned, similar to those described
in H. loosanoffi and canaliculata, were noted in a specimen of H. oculata from
Fipennies Ledge, Gulf of Maine.

The generally larger size of the northern individuals extends to the flagel-
lated chambers. In a specimen from Fipennies Ledge, Gulf of Maine, these range
from 36 x 26 to 43 x 33u. In a specimen from Block Island Sound, they range
in size from 21 x 21p to 26 x 20n.

DIsTRIBUTION IN NorTH AMERICA: Gulf of St. Lawrence to North Caro-
lina.

River and Gulf of St. Lawrence (Lambe, 1900b, p. 155; Whiteaves, 1901,
p. 15); coast of Nova Scotia (Lambe, 1896, p. 184; Whiteaves, 1901, p. 15); Minas

SYSTEMATIC STUDIES 61

FicurE 18. Haliclona oculata. Portion of skeleton in interior of branch, showing
extensive development of spongin. Spongin stippled. Long Island Sound (off Mans-
field Pt., East Haven, Conn.), 8 meters. YPM #810.

Ficure 19. Haliclona oculata. Portion of skeleton (section perpendicular to sur-
face) with spongin tracts. Spongin stippled. Fipennies Ledge, Gulf of Maine, 75
meters. YPM #995.

Figure 20. Haliclona oculata. Portions of skeleton from different parts of the
same colony. Channel Coast of France (near St. Valéry-en-Caux). YPM #1073. A
and B. Sections perpendicular to surface. C. Portion of skeleton from interior of
colony.

62 MARINE SPONGES

Basin, Bay of Fundy (Lambe, 1900b, p. 155; Whiteaves, 1901, p. 15; Hartman—
A. E. Verrill collections); Bay of Fundy, low water to 150 meters (Verrill, in:
Verrill and Smith, 1873, p. 742); off Kent Isl., N. B., 30 meters (Hartman); East-
port, Me. (Hartman—A. E. Verrill collections); Mount Desert Region, Me. (Proc-
ter, 1933, p. 93); Casco Bay, Me. (Verrill, 1874a, p. 44, 1874b, p: 364, and 1880,
p. 232; Kingsley, 1901, p. 161); Portland, Me., on wharf piles (Verrill, 1874a,
p. 133); Fipennies Ledge, Gulf of Maine, 80 meters (Hartman); Stellwagen Bank,
Massachusetts Bay, 40 meters (Hartman); Georges Bank (Smith and Harger, 1876,
p- 22; Hartman—A. E. Verrill collections); off Gay Head, Martha’s Vineyard,
8 to 30 meters (Verrill, 7n: Verrill and Smith, 1873, p. 742); Vineyard Sound, (Ver-
rill, 1880, p. 232, as Chalina oculata); Vineyard Sound, 2 to 16 meters (Verrill,
in: Verrill and Smith, 1873, p. 743, as Chalina arbuscula); Woods Hole, Mass.
(Allee, 1923, p. 175, as Chalina arbuscula; de Laubenfels, 1949, p. 9; Hartman;
see also the discussion by Sumner et al. 1913, p. 558); off Watch Hill, R. L. (Verrill,
in: Verrill and Smith, 1873, p. 742; also reported as Chalina arbuscula on p. J23);
Block Island Sound, 20—40 meters (Hartman); Long Island Sound, west to New
Haven, to 10 meters (Hartman); off Long Island, N. Y. (Rafinesque, 1819, p. 150,
as Spongia cespitosa); Great Egg Harbor, N. J. (Verrill, in: Verrill and Smith,
1873, p. 743, as Chalina arbuscula); Ft. Macon, N. C. (Coues and Yarrow, 1879,
p. 312, as C. arbuscula); North Carolina (Verrill, op. cit., p. 743, as C. arbuscula).

Haliclona loosanoffi sp. nov.

SHAPE AND SIZE OF COLONIES: Connecticut colonies (PI. 11) encrust rocks, shells,
and algae, and range in size up to 15 cm. in diameter and 1.0 to 1.5 cm. in thick-
ness. Vertical tubules are always present, varying in height from 2 to 33 mm.,
with basal diameters of 1 to 4 mm. The taller tubules often branch and anasto-
mose with neighboring ones. Sometimes the tubules bifurcate or trifurcate dis-
tally. Maryland colonies (Pl. 12, figs. 2, 3) range up to 2 to 3 cm. in thickness,
with short tubules (5 mm.) arising from the basal encrusting mass. The largest
colony from the Maryland population measures 9 x 7 cm. in horizontal dimen-
sions. Colonies with elongate branches are less frequent in the series from Mary-
land in the collections at Peabody Museum.

COoLor IN LIFE: Various shades of dark tan, gold, and drab (Maerz and Paul,
1950: PI. 11, F-5; Pl. 12, D-4; Pl. 13, F-5, F-6, F-7, G-7, I-7; Pl. 14, G-5, G-6, G-7,
H-6). Often the same colony will show gradations from a darker to a lighter
color; one such colony, collected in August, varied from gold (PI. 13, I-7) to beige
(PI. 12, D-4). The lighter colored portions of the colony seemed partially degener-
ate, possibly because of contact with mud beneath the rock on which the colony
was growing. Some of the larger colonies are tinged with pink or lavender.

Consistency: Soft and compressible.

SuRFACE: Sparsely hispid, resulting from the fact that multispicular tracts
frequently penetrate the surface, thus raising the dermal membrane into small
cones above the general surface of the colony. The surface tufts of spicules are
from 250 to 600u apart.

OscuLEs: Usually borne terminally on tubules; distributed as well along the
sides of the taller tubules. Circular in outline, with diameters ranging from 1.0
to 2.4mm; or elliptical, with diameters ranging from 0.8 x 1.5mm. to 1.5 x
2.0mm. (preserved specimens).

Pores: Distributed singly or in groups. Usually elliptical in outline, ranging
in diameters from 10 x 8p to 25 x 20u in preserved specimens.

SYSTEMATIC STUDIES 63

ECTOSOMAL ANATOMY (see fig. 21): The dermal membrane is thin and lacks
spicules; it is made up of a layer of exopinacocytes beneath which are one or two
layers of elongate amoebocytes. The dermal membrane is separated from the
endosome by extensive subdermal cavities, continuous with the incurrent and
excurrent channels. The dermis is supported by numerous mesenchymal tra-
beculae and vertical spicule tracts which traverse the subdermal cavities. ‘Ter-
minal spicules of the vertical tracts pierce the dermis at intervals of 250 to 600n.

ENDOSOMAL ANATOMY (see fig. 21): The endosome consists of trabeculae of
cells separated by the incurrent and excurrent canal systems. The flagellated
chambers are grouped in the trabeculae around excurrent canals; the canal sys-
tem is eurypylous. ‘The flagellated chambers are ellipsoidal in shape, with diam-
eters varying from 36 x 23u to 26 x 2lu. The choanocyte cell bodies are 2.3 to
3.0u in diameter; their nuclei are basal. The subdermal cavities as well as the
canals of the aquiferous system are lined by endopinacocytes. Cell strands, 15 to
40u in diameter and up to 850y in length, run through the endosome at inter-
vals. These strands are made up of numerous elongate cells containing basophilic
granules and occasional round cells with eosinophilic granules. Such cell strands
always have young spicules imbedded in them and may be compared to the
“cordons cellulaires” described by Herlant-Meewis (1949) in Spongilla. As Her-
lant-Meewis has suggested, these cell strands doubtless serve as auxiliary sup-
porting structures and also help to isolate and direct the spicules during their
formation.

SKELETON: Multispicular tracts joined by spongin (2 to 6 spicules per cross-
section) run vertically through the endosome, terminating in the dermal mem-
brane (fig. 22). Horizontally placed individual spicules connect adjacent tracts at
frequent intervals forming a loose network. The amount of spongin present varies
greatly from colony to colony and even in different parts of a single colony, rang-
ing from small amounts joining adjacent spicule ends to continuous fibers enclos-
ing spicules. Burton (1926) observed a similar range of variation in the British
haliclonid, Reneira cinerea.

The spicules are chiefly oxeas which taper gradually to sharp points and curve
gently at the mid-point. Some specimens have small numbers of styles and (or)
strongyles in addition to the oxeas (fig. 23). The oxeas vary greatly in size from
specimen to specimen, the range of mean values for 21 specimens (based on 100
measurements for each specimen) is: length, 85 to 156; width, 4.0 to 7.2. The
absolute ranges of spicule size for all 21 specimens are: length, 66 to 185u; width,
2.0 to 8.2u. Strongyles and styles are generally somewhat shorter than the mean
lengths of oxeas in any specimen.

GEMMULEs: Characteristic of this species is the formation of gemmules during
late summer (beginning in late August) and early fall. The gemmules form in a
basal layer in contact with the substratum to which they remain attached after
degeneration of the colonies (fig. 24; Pl. 12, figs. 1, 4). The gemmules are usually
hemispherical in shape, flattened on the lower surface. Occasionally they are
elongated into ellipsoidal or ovoid shapes. Dimensions of gemmules from Connec-
ticut specimens: horizontal diameter, 315 to 515y; height, 215 to 330u. Gemmules
of the Maryland colonies examined are larger, ranging in horizontal dimensions
from 360 x 325u to 1000 x 725u.

In life the gemmules are white in color in Connecticut colonies and are difh-
cult to detect on oyster or barnacle shells. The gemmules of Maryland colonies,

MARINE SPONGES

64

‘96281 WdA ‘UWUOD “asseuowweyy ‘(aoezIns 0} IepNIIpuadiad uonsas) YJouvsoo] vuojI11vF] Jo AWoyeuy *[Z zANIIT
Sajnoids pepnyjou

A001
YIM ps0o [199 ee

s3}A90qa0wD
jOWAYoUasow

JQUUDYD JUd4aJJD

JauUDYD judaJasje

AyIADo suDIquow
}OW4epqns jOwJsep

SYSTEMATIC STUDIES 65

20p

FicurE 22. Portion of skeleton of Haliclona loosanoffi (section perpendicular to
surface). Milford Harbor, Conn. YPM +859B.

FicurE 23. Spicules of Haliclona loosanoffi. Oxeas, styles, strongyles. Pine Orchard
(Branford), Conn. YPM +1833.

preserved in alcohol, are yellow. Each gemmule is enclosed in a spongin capsule in
which special oxeas are imbedded. The base has relatively few spicules, however.
The gemmule spicules are slightly shorter and wider than the skeletal spicules;
they tend to be straight or only slightly curved. The range of mean spicule size of
four specimens (25 measurements per specimen) is: length, 83 to 954; width, 4.1
to 6.3u. Absolute range in size: length, 69 to 1024; width, 3.3 to 6.9u.

In early stages of formation, the gemmules consist of elongate cells, presum-
ably archaeocytes, containing minute, sparsely distributed granules. It appears in
sections that these cells migrate to the base of the colony from throughout the en-
dosome. The engulfing of trophocytes by archaeocytes, as described by Leveaux
(1939) in spongillids, was not observed. The basal layer of spongin is laid down
first, and contains few spicules. Later, the entire mass of cells becomes enclosed in
a spongin sheath in which oxeas are secreted (fig. 25). A layer of granule-free cells
underlies the spongin sheath, and it is presumed that these secrete the spongin. In
mature gemmules the granules of the interior cells have increased greatly in num-
ber and size (up to 2.5u in diameter) and many appear to have broken out of the
cells (possibly an artifact). Preliminary histochemical studies'® of these reserve

16 Periodic acid-Schiff test for polysaccharides, negative; Millon reaction for proteins,
weakly positive; Sudan Black B test for lipids, positive. ‘The granules stain black with
iron haematoxylin and red with Mallory triple. All staining was done on Bouin-fixed
material, preserved in 70 per cent alcohol for several years. Conclusive results must await
further studies with fresh material.

66 MARINE SPONGES

Ce ake
Vie SUSE va 0 ate
CMDS ATA

BOX: Hs I GF <p

Dee cose
WOE SO Paes
O UF ey Dyce

Ficure 25. Cross-sections through gemmules of Haliclona loosanoffi. Left: mature
gemmule covered by spicule-laden membrane of spongin. Right: Immature gem-
mule before formation of the spongin membrane. Pine Orchard (Branford), Conn.
YPM 31832F.

granules suggest that they contain lipids and proteins, thus resembling the reserve
substances in spongillids (Pourbaix, 1935).

Eccs AND EMBRYOS: Eggs and embryos were present in specimens collected in
late August and early September. Eggs alone were found in colonies gathered in
late September and early October. The eggs and embryos are located throughout
the endosome in the cellular trabeculae. They appear in sections to occur in cavi-
ties, and the embryos are enclosed in limiting membranes (spongin?) adhering to

SYSTEMATIC STUDIES 67

the outside of which is a layer of elongate cells. The cells of the embryos are filled
with basophilic granules, presumably food reserves. Late stages of the embryos
contain numerous thin oxeas arranged longitudinally throughout the interior.
Dimensions: eggs, spherical to ovoid in shape, with diameters ranging from 25 x
25u to 43 x 35u; nuclear diameters, 8 to 15; nucleolar diameters, 3.5 to 5.0u;
embryos, subspherical to ellipsoidal in shape, with diameters ranging from 150 x
130 to 215 x 185y. The larval oxeas vary in size from 45 x 1.5 to 55 x 1.5u.
Free-swimming larvae have not been found as yet.

HototyPe: Yale Peabody Museum No. 614. Colony with gemmules collected
below mean low water at Milford Harbor, Connecticut, in October, 1947.

REPOSITORIES OF OTHER TYPE MATERIAL: U. S. National Museum, Museum of
Comparative Zoology, British Museum (Natural History). About 35 lots of speci-
mens from six localities in central Connecticut and 12 lots from Solomons Isl.,
Maryland, were studied.

Type LocaLity: Milford Harbor, Connecticut, from mean low water to a
depth of two meters. Common as a fouling organism on submerged piles, floats,
and shells.

FURTHER DISTRIBUTION: Long Island Sound to Maryland. This is a common
sponge on the coast of central Connecticut, where it has been collected at Ham-
monasset State Park, Pine Orchard (Branford), Double Beach (Branford), Light-
house Pt. (New Haven), and Bradley Pt. (West Haven), as well as at Milford. It
usually encrusts the undersides of large rocks at and below mean low water level,
occasional colonies occurring in tide pools up to mid-tide level. It is also a fouling
organism on wharf piles, floats, and shells, chiefly in places not subject to direct
sunlight. Several specimens have also been collected on oyster shells off Wood-
mont, Connecticut, at a depth of ten meters. Dr. I. M. Newell collected about 12
colonies of the sponge from wharf pilings at Solomons Island, Maryland. A simi-
lar sponge was found growing under the rocks of a breakwater at Stone Harbor,
N. J. by the writer but the colonies lack the diagnostic gemmules, and their iden-
tity is still uncertain.

The numbers of this sponge show large annual fluctuations in some localities.
In 1947 and 1948, it was a common fouling organism at Milford; in 1953 and
1954 not a single colony was found at this locality. In 1953 it was abundant at
Hammonasset State Park; in 1954 only a few colonies could be found there.
Nevertheless, the sponge was found in abundance at Pine Orchard in 1953 and at
Double Beach in 1954, indicating that the population decline was not general in
the region.

VARIATION: During the course of the studies of this species, great variations
were noted in spicule size, and it seemed of interest to determine whether this
character is correlated with geographic range and whether several sibling species
might be present. Spicule slides of samples of 22 colonies from five localities were
prepared, and length and width measurements were made of 100 spicules on each
slide. The samples used for preparing the slides were about 0.5 cm? in size and
were chosen to include all levels of the sponge. The data are given in Table 21.

The coefficient of variability of the mean values for spicule length of the 22
colonies is very high (C.V.=16.8), suggesting that the samples are not homogen-
eous. If the coefficient of variability of samples from each of the five localities is
considered separately, those of Hammonasset and Double Beach are seen to be
especially high (Table 20).

These data suggest the possibility that two sibling species of Haliclona are

68 MARINE SPONGES

TABLE 20
COEFFICIENT OF VARIABILITY OF SPICULE SIZE IN HALICLONA
LOOSA NOFFI
LOCALITY NO. OF MEAN SPICULE Cave
COLONIES LENGTH
Hammonasset 8 104 + 16.2 15.6
Double Beach 5 127 + 18.4 14.5
Milford 4 100+ 4.3 4.3
Pine Orchard 4 96+ 8.0 8:3
Lighthouse Pt. 1 97 Insufficient data
First two localities 13 113 + 20.4 18.1
Last three localities 9 98+ 6.4 6.5
ALL LOCALITIES 22 107 + 17.8 16.8 ‘

present on the Connecticut coast, one, with shorter spicules being found at all lo-
calities listed, the other with longer spicules, occurring along with the former at
Double Beach and Hammonasset. If the mean spicule lengths of the Hammonas-
set and Double Beach colonies are compared with those of Milford, Lighthouse
Point, and Pine Orchard colonies considered together, P=slightly>0.05, very
near the borderline of significance.

If two species with differing spicule dimensions are actually present in the
Hammonasset and Double Beach populations, other characters would be expected
to show a correlation with spicule size. Such has not been found to be the case,
so far. Tubule height and the occurrence of gemmules, e.g., show no correlation
with spicule size. On the basis of present evidence, the several populations of
Haliclona loosanoffi may be considered as representatives of a single species,
highly variable in respect to spicule size and colony form.

OTHER SPECIES TO BE COMPARED WITH Haliclona loosanoffi: Encrusting haliclon-
ids are notoriously difficult to define on the species level. They are often highly
variable in regard to external form, skeletal characteristics, and color, so that
these characters are reliable only if large samples of a population are studied. In
the past many species have been described on the basis of one or two specimens,
thus complicating the question of synonymy in the genus. The validity of the
new species, Haliclona loosanoffi, is based most firmly on the occurrence of gem-
mules, although spicule dimensions and color also assume a significance when
large samples are considered. It is probable that studies of larvae will also be im-
portant in distinguishing haliclonids when enough is known about them, just as
they have been useful in the difficult genera, Halichondria (Topsent, 1911) and
Halisarca (Lévi, 1953a, 1956).

Haliclona loosanoffi is superficially similar to H. permollis (Bowerbank), a

SYSTEMATIC STUDIES

TABLE 21

SPICULE DIMENSIONS OF HALICLONA LOOSANOFFI*

LOCALITY AND YPM
CATALOGUE NUMBER

Hammonasset, Conn.
No. 1821

Hammonasset, Conn.
No. 1823

Hammonasset, Conn.
No. 1825

Hammonasset, Conn.
No. 1826

Hammonasset, Conn.

No. 850

Hammonasset, Conn.
No. 1822

Hammonasset, Conn.
No. 1824

Hammonasset, Conn.
No. 1829

Pine Orchard (Branford),

Conn. No. 1832

Pine Orchard (Branford),

Conn. No. 1833

Pine Orchard (Branford),

Conn. No. 1834

Pine Orchard (Branford),

Conn. No. 827

SPICULE LENGTH
RANGE AND MEAN

76-120-149p

119-138-152

78-88-98

70—91-107u

82—106-131p

90-104-115y

82-93-1003

70-92-1111

69-85-96

70—100-115

7892-103

94-106-115p

Je

4.

ANS

SPICULE WIDTH
RANGE AND MEAN

3-5.0-7.3p

1-5 .9-7.8u

.7-4.4-6. 2p

.1-4.2-4. 9p

lose 2

3-4. 3-5 3

.1-4.0-4.5p

9-4.2-4.9u

.3-6. 1-7 .3p

.3-5 .6-8.2u

7-5 9-8. 2m

M27 2282p

* All measurements based on 100 spicules per specimen.

69
REMARKS
Style:
102 x 4u
Styles:
86-lllu x 4.9-8.2u
Strongyles:

94-111p x 6.2-8.2u

Style:

83 x 6.94
Strongyle:
63 x 7.3

Strongyles:

94-107u x 6.2-8.2u

Strongyle:
70x 5.3u

(Continued)

cosmopolitan species according to de Laubenfels (1936, 1949, etc.). The mean spic-
ule dimensions given by the latter author (1949, see Table 22) are considerably
larger than those of the Connecticut population. However, the spicule lengths
given by Bowerbank in his description (1866) and illustration (1874) of the type

70 MARINE SPONGES

TABLE 21—Concluded
SPICULE DIMENSIONS OF HALICLONA LOOSA NOFFI*

LOCALITY AND YPM SPICULE LENGTH SPICULE WIDTH REMARKS
CATALOGUE NUMBER RANGE AND MEAN RANGE AND MEAN
Double Beach (Branford), 83-133-152u 2.0—5 .0-6.6u
Conn. No. 1838
Double Beach (Branford), | 96-131-158u 3.3-5 .8-6.9u Styles:
Conn. No. 1848 119-219 x 5.0-6.3u
Double Beach (Branford), | 103-156-185u 4.1-5.6-7.0Ou
Conn. No. 1956 D-1
Double Beach (Branford), | 86-112-144u 4.1-5.8-7.8y
Conn. No. 1956 E-1
Double Beach (Branford), | 86-103-152u 4.1-5.9-7.8u
Conn. No. 1956 E-2
Lighthouse Point, New 66-97-112u 2.3-6.0-7.3u Styles:
Haven, Conn. No. 841 89-99u x 5.9-6.6pu
Milford Harbor, Con- 73-101-116p 2.6-5.3-6.6u
necticut No. 859
Milford Harbor, Con- 90-103-115u AA 7 1-8
necticut No. 614
Milford Harbor, Con- 90-104-115 2147-6 6p
necticut No. 2139
Milford Harbor, Con- 78-93-107u 2.5-3.8-4. 5p
necticut No. 1876
Solomons Isl., Md. 78-104-119p 2.9-6 .3-7.8u
No. 623
Solomons Isl., Md. 90-111-127y 4.1-6.2-8.2u
No. 627

* All measurements based on 100 spicules per specimen.

are comparable to those of the Connecticut forms. De Laubenfels (1949) describes
the color of living colonies of H. permollis as “‘a very distinctive lavender”; dead
or “pathological” specimens are “dull, pale brown.” Lavender-tinged specimens
of H. loosanoffi are the exception, most colonies being tan to golden brown, in
Connecticut, at least. In skeletal architecture the two species are very similar.
However, gemmules have not been reported from H. permollis; it seems unlikely
that they would have been overlooked by all of the many authors who have stud-

SYSTEMATIC STUDIES 71

ied this species. The present author concludes that H. loosanoffi may be distin-
guished from H. permollis as a gemmuliferous species with smaller spicules and
color differences.

Another species to be compared with H. loosanoffi is Reniera tubifera George
and. Wilson (1919), found at Beaufort, N.C. De Laubenfels (1947) considered this
species as a synonym of Haliclona permollis, but in so doing he overlooked two
important characters mentioned by George and Wilson concerning the consis-
tency and dermal structure of their species. These authors state that Reniera tubi-
fera is “not soft, but quite fragile,” and they describe and figure “a distinct uni-
spicular reticulum” in the dermis. The present writer has studied a series of
haliclonid-like sponges collected at Beaufort Harbor, N.C., by Dr. Irwin M. New-
ell for the Peabody Museum. It is immediately apparent from consistency alone
that two species are present in the series. One is soft and compressible; the other
is “not soft, but quite fragile.” Indeed, the colonies of the latter species, preserved
in alcohol, have broken into many pieces just through handling of their con-
tainer. The spicules of the first species are small; those of the second, large (see
Table 22). The dermis of the first lacks spicules; that of the second is provided
with “a distinct unispicular reticulum.” There is no question that the second spe-
cies in Newell’s collection is Reniera tubifera, but the dermal spiculation suggests
that it is not a Haliclona but rather a member of the genus Adocza, following the
concept of that genus discussed by Burton (1934) and de Laubenfels (1936). ‘The
first-mentioned species is clearly a Haliclona; although it agrees with H. loosan-
offi in spicule size and skeletal arrangement, further study is needed to identify
it specifically.

Several other haliclonids recorded from the Atlantic Coast of North America
may be compared with H. loosanoffi. Reniera mollis Lambe, 1893, recorded by its
author from Labrador and the Baie des Chaleurs (1896), from Hudson Bay
(1900a), and from Davis Strait and Hudson Strait (1900b) is an offshore species
with large spicules and pores and is massive to lobate in form. Reniera heteroft-
brosa Lundbeck, 1902, recorded by Procter (1933) from the Mt. Desert region,
Maine, has large spicules arranged in an irregular manner, resembling Halichon-
dria. Haliclona palmata (Ellis and Solander, 1786) described by de Laubenfels
(1949) from Woods Hole, has very small spicules enclosed in prominent spongin
fibers.

Tubule-bearing colonies (Pl. 11, figs. 12 through 16) of Haliclona loosanoffi
show a striking resemblance to the European species, Halichondria montagutt
Fleming, 1828, as figured by Johnston (1842, Pl. 6, fig. 1). Other colonies (PI. 11,
figs. 8, 9, 10) resemble figures of Chalina montagui (Bowerbank, 1866) and Cha-
lina flemingit Bowerbank, 1866, given by Bowerbank (1874, Pl. 68, figs. 1-5).
American specimens fit the descriptions given by these authors based largely on
external form, color, and skeletal characteristics. Spicule dimensions recorded by
Bowerbank for montagut and flemingit (based on two spicules in each case) fall
largely within the range of spicule sizes observed in Haliclona loosanoffi. Unfor-
tunately it has not been possible to study Bowerbank’s type material nor to obtain
fresh material from England. That the American species is distinct from the Brit-
ish ones is supported by the disjunct pattern of distribution. It is unlikely that
Haliclona loosanoffi occurs north of Cape Cod as would be expected in the case
of a species occurring on both the British and American coasts. The author has
made a thorough survey of the intertidal sponge fauna at six localities north of
Cape Cod (Cape Ann, Mass.; Kittery, Maine; Cape Elizabeth, Maine; Boothbay

72 MARINE SPONGES

TABER 22
COMPARISON OF AMERICAN AND BRITISH HALICLONIDS

SPECIES AND LOCALITY AUTHOR SPICULE REMARKS
DIMENSIONS
Haliclona loosanoffi Hartman Connecticut: Pores: 10 x 8u to
Connecticut to Maryland 66-107-185y 25 x 20u
x 2.0-5.4-8.2u | Oscules: 1.0 to 2.4 mm.
Maryland:
78-108-127y
x 2.9-6.3-8.2pu
Haliclona sp. Hartman 111-752-180
New Jersey x 2.9-5.5-7.8u
Haliclona sp. Hartman 78-99-1194
Beaufort, N. C. x 2.1-4.9-7.4y
Haliclona permollis de Laubenfels, 130-150-170u Oscules: 1 to 5 mm.
Woods Hole 1949 x 6-7-8
Haliclona palmata de Laubenfels, 55—60u x 2-34 | Oscules: 1 to 3 mm.
Woods Hole 1949 Fibers: up to 40u
Reniera heterofibrosa Procter, 1933 121-162 (mode)
Mt. Desert Region, —203u
Maine
Reniera mollis Lambe, 1896 170-299 Pores: 65u
Northeastern Canada x 6-9u Oscules: 5 mm.
Isodictya permollis Bowerbank, 113u; 116y
Great Britain 1866; 1874
Chalina montagutii Bowerbank, 116u x 8.5u
Great Britain 1866; 1874 127u x 6.6u
Chalina flemingit Bowerbank, 85u x 3.4u
Great Britain 1866; 1874 102u x 3.8u
Reniera tubifera George and 125-170u Pores: 50u
Beaufort, N. C. Wilson, 1919 x 3-8u Fibers: 30-100
Adocia tubifera Hartman 131-159-176pu
Beaufort, N. C. x 2.9-7 .4-9 .Ou

Harbor, Maine; Rockland, Maine; and Kent Island, N. B.) and has not found this
haliclonid.

The species is named in honor of Dr. Victor L. Loosanoff, Director of the
Biological Laboratory of the U. S. Fish and Wildlife Service, Milford, Connecticut.

SYSTEMATIC STUDIES 73.

Haliclona canaliculata sp. nov.

SHAPE AND SIZE OF COLONIES: The colonies form thin encrustations on rocks,
shells, and algal holdfasts, ranging in size up to 10 cm. in diameter and from 2 to
8 mm. high (PI. 15, fig. 1).

CoLor IN LIFE: Winter specimens: various shades of light buff, maple, and
light tan (Maerz and Paul, 1950: Pl. 11, D-4, G-6; Pl. 12, B-4, C-6, D-5; Pl. 13,
D-6). Summer specimens: dark tan to brown, drab, and gold. (PI. 12, B-4; Pl. 13,
Doh, F-7; Pl. 14, D-5, E-4; 1-7).

ConsIsTENCy: Moderately soft.

SuRFACE: The surface of the colonies is raised at intervals into pointed,
rounded, or ridge-like projections, 1-3 mm. high, where spicule tracts and cellular
trabeculae traverse the subdermal cavities and reach the dermal membrane, push-
ing it above the general surface of the colony. Radiating subdermal excurrent
channels converge upon the oscules and provide the surface with stellate patterns.
These channels are often separated and rendered more prominent to the naked
eye by ridges of choanosome which rise above the general surface of the colony.
The circular openings of the incurrent channels leading from the subdermal cavi-
ties into the choanosome can readily be seen through the dermis by the unaided
eye; these openings give the colony a punctate appearance (PI. 5, fig. 2).

OscuLEs: Circular or elliptical in outline, ranging in diameter from 0.8 x 0.6
mm. to 1.8 x 1.4 mm. in preserved specimens. The oscules are not elevated above
the general surface of the colony as a rule; occasionally they are located on the
summits of low mounds.

Pores: Occurring in groups above subdermal cavities, elliptical in shape, rang-
ing in diameter from 30 x 16u to 57 x 50u.

EcTosoMAL ANATOMY: The thin dermal membrane is composed of a layer of
exopinacocytes underlain by one or two layers of elongate amoebocytes. The
groups of dermal pores open into extensive subdermal cavities which in turn lead
into wide incurrent canals. The openings into the latter from the subdermal cavi-
ties are readily apparent through the thin dermal membrane and appear to the
naked eye like pores in the surface of the colony as noted above. Thin trabeculae
of mesenchyme as well as occasional spicule tracts run through the subdermal cavi-
ties to the dermis, the spicules piercing the surface at intervals.

ENDOSOMAL ANATOMY: The endosomal mesenchyme consists of membranes or
trabeculae of cells bounded by the extensive system of incurrent and excurrent
canals. The canal system is eurypylous, the flagellated chambers opening directly
into the excurrent canals. The flagellated chambers are ellipsoidal in shape, with
diameters varying from 25 x 20u to 35 x 25u. The choanocytes range in diameter
from 3.0 to 4.0u. The excurrent canals of the endosome open into wide subdermal
cavities which lead to the oscules. Both incurrent and excurrent canals are lined
with endopinacocytes.

As in Haliclona loosanoffi, strands of elongate amoebocytes are frequent in the
endosome. Tracts of spicules, free from spongin, lie in these cell strands which
are 45 to 90u in width and run chiefly parallel to the surface of the colony.

SKELETON: The basic framework of the skeleton consists of vertical multispicu-
lar tracts (three to ten spicules per cross section) held together by small quantities
of spongin. The vertical tracts are joined by horizontally placed individual spic-
ules or short tracts; in addition there are many irregularly arranged spicules be-
tween the tracts (fig. 26).

74 MARINE SPONGES

M4
20p
FicureE 26. Portion of skeleton of Haliclona canaliculata (section perpendicular

to surface). Spongin stippled. Dermis shown as dashed line. Double Beach (Bran-
ford), Conn. YPM 42016B.

Figure 27. Spicules of Haliclona canaliculata. Oxeas, styles, strongyles, centrotylote
oxea on right. Double Beach (Branford), Conn. YPM #1955D.

Long horizontal multispicular tracts are also characteristic of this species;
these are free of spongin but are associated with strands of cells as described
above. Such tracts have been traced for distances up to 3mm. in sections perpen-
dicular to the surface; they are about 75u thick.

The spicules are oxeas, straight or gently curved in the middle, tapering grad-
ually to sharp points (fig. 27). In some cases there is a step-wise reduction in width

SYSTEMATIC STUDIES

TABLE 23

SPICULE DIMENSIONS OF HALICLONA CANALICULATA*

LOCALITY AND

YPM CAT. NO.

Hammonasset
1827

Double Beach
1955D

Double Beach

1955E-1

Double Beach

1955E-2

Hammonasset
854

Double Beach

1785

Double Beach

1788

Double Beach
1791

Double Beach

1803

Double Beach

1811

Double Beach
1813

Double Beach

1816

Double Beach

1874

All summer
specimens (5)

All winter
specimens (8)

DATE OF
COLLECTIONS

August

September

September

September

October

January

January

January

February

February

February

February

March

August to
October

January to

March

RANGE AND MEAN OF
SPICULE LENGTH X WIDTH

86-102-115u
x 3.3-4.3-6. 2p

107—125-139y
x 4.1-5.9-7.4u

98-125-148u
x 4.5-6.7-7.8u

94-113-123u
x 4.1-5.3-7.0u

82—112-135u
x 3.7-6.1-8.2u

103-116-131
x 4.1-7.9-8.2u

103-122-135
x 5.3-6.9-8.2y

904-112-127
x 5-3-70-8) 2p

74-109-131u
x 5.3-7.7-8.2u

98-113-127u
x 4.5-/.2-8.2u

94-119-131p
x 4.1-5.7-8.2p

98-113-123u
x 5.3-7.4-8.2u

98-118-135
x 4.9-7 8-8 .2u

82-115-148u
x 3.3-5.7-8.2u

74-115-135u
x 41-728. 2p

* All measurements based on 100 spicules per specimen.

REMARKS

Strongyles:
78-107 x 5.3-8.2u

Styles:

103-107u x 5.3-6.1p

"6 MARINE SPONGES

at the ends. Styles and strongyles in small numbers are found in many specimens
along with the dominant oxeas. There is less variation in the lengths of spicules
than was observed in Haliclona loosanoffi. The range of mean spicule sizes for 13
specimens (based on 100 measurements per specimen) is 102 to 125y in length
and 4.3 to 7.9u in width. The absolute range in length is 74 to 148; in width, 3.3
to 8.2u. Styles and strongyles are shorter than oxeas. The coefficient of variation in
spicule length is 5.8; in width, 16.7.

HototyrE: YPM No. 2017. Collected on the underside of a rock, 0.5 feet be-
low mean low water; Double Beach (Branford), Connecticut, Sept. 17, 1955.

REPOSITORIES OF OTHER TYPE MATERIAL: U. S. National Museum; Museum of
Comparative Zoology, Harvard; and British Museum (Natural History). About
20 lots were studied.

Type Locatity: Double Beach (Branford), Conn. Common under rocks at
mean low water and below.

FURTHER DISTRIBUTION: Two specimens were collected at Hammonasset State
Park (Meig’s Point), Connecticut.

Discussion: Haliclona canaliculata differs from the other common encrusting
haliclonid of Long Island Sound, H. loosanoffi, in being a perennial species. Colo-
nies undergo some degeneration with the onset of cold weather, but portions of
each colony remain alive throughout the winter months. ‘These colonies show sev-
eral differences from summer colonies: (1) they lack choanocytes, the endosome
consisting of aggregates of amoebocytes (2) they possess thicker spicules. A more de-
tailed investigation of the morphology and feeding habits of winter colonies is in
progress. The occurrence of thicker spicules in winter colonies may reflect the
greater availability of silicates during the winter, a fact observed in many hydro-
graphic studies (Sverdrup, Johnson, and Fleming, 1946). This suggestion is sup-
ported by the experimental work of Jérgensen (1944) who found that the thick-
ness (but not the length) of the microscleres of Spongilla lacustris increases with
increasing silica content of the medium.

COMPARISON WITH OTHER SPECIES: In external form this species differs from
other encrusting haliclonids of the American coast in the absence of branches and
raised oscules. Conspicuous features of living and well-preserved colonies are the
subdermal excurrent channels which appear to radiate out from the oscula, and
the openings from the subdermal incurrent cavities into the incurrent endosomal
canals which look like pores in the surface of the colony. ‘The spicules are smaller
than those of most other encrusting haliclonids (see Tables 22 and 23) of the
American coast, except Haliclona loosanoffi, from which it differs in external
shape and in lacking gemmules.

Reniera plana Topsent (1892) is a flat encrusting haliclonid, warranting com-
parison with H. canaliculata. The Mediterranean sponge has its spicules arranged
in a unispicular network, however, and the spicules are much larger (235-250u x
9) than those of the species described herein.

SUMMARY

1. The systematics of twelve species of Demospongiae from Block Island
Sound and Long Island Sound is considered. Of these, two are described as new:
Haliclona loosanoffi and Haliclona canaliculata. The taxonomic status of Verrill’s
species, “Chalina arbuscula,” and of de Laubenfels’ species, “Neospertopsis deich-
manni,”’ is discussed.

2. Trends of variation of the skeleton of monaxonid sponges correlated with

SYSTEMATIC STUDIES fi!

latitude are exemplified. Megasclere size is shown to increase in northern regions
in Haliclona oculata; styles are replaced by oxeas in the more northern of two
species of Isodictya studied.

3. Of the twelve species considered in this work, three (Cliona celata, Cliona
lobata, and Cliona vastifica) are apparently cosmopolitan in distribution. Suber-
ites ficus is widely distributed in the North Pacific and North Atlantic, reaching
the Mediterranean Sea and the offshore waters of northwestern Africa in the lat-
ter region. Haliclona oculata is found in waters of the continental shelf of Europe
south to Portugal and of North America south to North Carolina.

Isodictya deichmannae and Microciona prolifera are both widely distributed
along the North American Coast, the latter species having been reported from
the Mediterranean area as well, but this requires verification. Block Island
Sound is the southernmost recorded occurrence of I. deichmannae. Halichondria
bowerbanki is found on the American Coast from the southern shores of Cape
Cod to Long Island Sound and also on the coasts of England and France.

The northernmost recorded occurrence of both Cliona truitti and Haliclona
loosanoffi is Long Island Sound. Haliclona canaliculata is known at present only
from Long Island Sound.

Lissodendoryx isodictyalis is widely distributed in tropical, subtropical, and
warm temperate seas. Its occurrence on the Atlantic Coast of North America as
far north as Woods Hole is reported here.

II, LIFE HISTORY STUDIES
PREVIOUS WORK

Detailed studies of the life histories of sponges especially in regard to the sea-
son of larval attachment, survival after attachment, and winter survival are not-
ably absent from the literature of this group of organisms. Such data as are re-
corded have usually been incidental to embryological work or to general studies
of fouling organisms. Topsent (1887), however, has recorded the breeding sea-
sons of the common sponges at Luc, on the Channel Coast of France, and Lévi
(1956) has given a series of records for Roscoff, France.

The studies of Coe (1932), Coe and Allen (1937), and McDougall (1943) give
some attention to the common sponges occurring in the regions of their work,
and these will be summarized here.

Coe (1932) found two sponges settling on his experimental blocks at La Jolla,
California. Concerning one, a simple calcareous sponge identified as a species of
Grantia (in a later paper determined as Rhabdodermella nuttingt Urban), Coe
observed that, whereas growth occurs throughout the winter and spring, repro-
duction is limited to the warmer months. An unnamed white encrusting sponge
[later identified as Leucetta losangelensis (de L.)] is said to grow to a diameter of
six inches or more from the first week in September to the following July. Pre-
sumably larvae of this species settle in late summer. In 1937 Coe and Allen re-
viewed the studies on the growth of sedentary marine organisms which had been
carried on for nine successive years at the pier of the Scripps Institution of Ocean-
ography at La Jolla. Rhabdodermella nuttingi Urban and Leucetta losangelensis
(de L.) were described as characteristic organisms of the summer months but no
precise dates of settling were indicated. It was noted that sponges are more com-
mon on blocks which have remained in the water for several months than on
freshly submerged surfaces. Oyster shells were found to be especially favorable
for larval sponge attachment.

McDougall (1943) made a careful study of the pile-dwelling invertebrates at
Beaufort, North Carolina, and reported on two species of sponges, Reniera tubt-
fera George and Wilson (—Adocia tubifera) and Microciona prolifera (Ellis and
Solander). The settling period of Adocia tubifera was found to last only about
thirty days in June and July after which the colonies spread out over the tiles,
often smothering low-growing forms such as barnacles and tubicolous annelids as
well as encrusting the stalks of hydroids. Microciona prolifera was only occasion-
ally found on the experimental tiles although it is a common sponge in the region.
Attached larvae were first noticed on August 12 and probably continued to set
throughout September. Growth was most rapid and extensive on tiles just above
the mud line, the finger-like processes characteristic of the species appearing on
these colonies in mid-December. Although McDougall found that most species of
sponges grow most vigorously during the spring months at Beaufort, he observed
that Adocia tubifera grows best in summer and Microciona prolifera in winter.

McDougall discovered that current has a pronounced effect on the settling of
larvae of Adocia tubifera. He devised a current box consisting of five chambers in
each of which water flowed at a different velocity and found a predominance of
settling in currents of intermediate velocity (<0.5 knot) for the Beaufort area.

78

LIFE HISTORY STUDIES 79

No significant effect of light on the settling of Adocia tubifera larvae was observed
by McDougall.

Pomerat and Reiner (1942) found a “small white sponge” growing on the
plates which they submerged at Pensacola, Florida, in an investigation of the ef-
fects of light and surface angle on attachment of larvae. Unfortunately, they gave
no information about the influence of these factors on the settling of the sponge,
as well as no clue as to its identification.

A number of investigators have recorded life history notes about calcareous
sponges incidental to embryological studies. Dendy (1914) found that Grantia
compressa (Fabr.) is an annual sponge at Plymouth, England, where it grows
most rapidly in the winter and spring and degenerates in autumn. The breeding
season begins in early April when germ cells are abundant; mature embryos were
observed leaving the oscules of sponges in June. There were also many advanced
stages of embryos observed in August, although germ cells were no longer com-
mon at this time. According to Dendy, Orton has observed two breeding seasons
for Grantia compressa at Plymouth. In June embryos are discharged from large
specimens of the previous season, these colonies subsequently degenerating. ‘The
June embryos give rise to colonies which, though still very small, produce numer-
ous embryos in October. Thus the same colonies probably breed twice during
their life histories, once in late autumn and again in late spring. Jorgensen (1917,
1918) made a study of the reproductive cycle of the same species at Cullercoats,
Northumberland. By examining colonies of this species every few weeks over the
period of a year she was able to demonstrate that only one breeding season exists
for Grantia compressa on the Northumberland coast. Small germ cells were first
detected in June. Specimens examined in July contained both eggs and embryos;
by August 17 the eggs were greatly reduced in number, but late-stage embryos
were still common. No embryos were found later than early September. In these
more northern waters, the colonies of the year do not seem to produce embryos
in late fall as Orton observed at Plymouth.

Duboscq and Tuzet (1937) stated that all stages in the development of eggs
and embryos were present in colonies of Grantia compressa in July at Roscoff, on
the Channel Coast of France. They reported that in Sycon raphanus at Banyuls-
sur-mer, on the other hand, the various developmental stages of germ cells and
embryos tend to occur simultaneously in any one colony so that only one stage
is regularly found at a time; if two stages are present, they are widely separated
in age. The breeding season in this species continues from February through No-
vember.

Several authors have recorded the breeding seasons of the clionids. Grant
(1826b) noted that colonies of Cliona celata were filled with eggs in March and
April along the coasts of Scotland. ‘Topsent (1887; 1900) has reported that speci-
mens of this species from the Channel Coast of France are filled with eggs in the
fall (late September and October). In the Mediterranean region reproduction
must be earlier (probably in mid- or late summer) than in the English Channel,
for Topsent (1900) found no trace of eggs in specimens from Banyuls-sur-mer
during the period from October to March. Grant’s observation is difficult to cor-
relate with the more southern records which suggest that the formation of repro-
ductive cells is initiated by high temperatures. As will be seen later, a similar cor-
relation of breeding with temperature is also found in the western North Atlantic.

Concerning Cliona vastifica, Topsent (1887; 1900) has reported that eggs are
abundant in the colonies in late September and October in the English Channel.

80 MARINE SPONGES

Nasonov (1883, 1924) found egg-filled colonies of Cliona stationis (=C. vastifica ?)
in the Black Sea in June and July. Again the implication is that high tempera-
tures, such as occur in mid- or late summer on the Channel coasts of France, and
and earlier in the Mediterranean region, initiate the reproductive cycle.

Tuzet (1930) found eggs in Cliona viridis at Banyuls-sur-mer from July
through September. Volz (1939) found eggs in colonies of Cliothosa hancockti
(Topsent) at the end of April; a larva of Thoosa mollis Volz was observed in
July. Both of these records are from Rovigno.

The observations on clionids summarized here suggest that these sponges be-
gin their reproductive cycle when the water is warmest in temperate regions; late
summer and early autumn are the seasons of egg and larva production on the
coasts of northern France, in Britain, and in New England. In warm temperate
regions the reproductive cycle begins earlier in the summer or even in late spring
and generally lasts longer.

Topsent (1887; 1911) observed a difference in the periods of larval production
in two species of Halichondria found along the Channel coasts of France. He
found that the larvae of H. panicea are released in May and early June, whereas
those of H. bowerbanki (= coalita) are freed in August and September.

Additional records of the breeding seasons of sponges are found in Table 24.
(See also Lévi, 1956, pp. 8-9.)

METHODS

In the present study observations were made on the period of larval settling
of the sponges of Milford Harbor, Connecticut, during the summers of 1947 and
1948, by suspending wire trays or bags filled with mollusc shells and rocks from a
floating raft anchored in the harbor or from the dock of the Milford Biological
Laboratory.

During the summer of 1947 the investigations were directed toward finding
out whether boring sponges (genus Cliona) show any preference for particular
species of molluscs and whether they settle preferentially on living molluscs or
on dead shells. To this end an assortment of molluscs was planted out as listed
in Table 25.

The molluscs were placed in a large wire tray (6 ft. x 3 ft. x 6 in.) which was
suspended from a floating raft anchored about 50 yards from shore in water six
feet deep at the mean low level. (The mean range of tides in Milford Harbor is
6.6 feet.) The tray was hanging about 314 feet below the surface of the water so
that the distance from the bottom varied from a mean of 21% feet to a mean of
nine feet, depending upon the tides. The water in which the raft was anchored
was exposed to full sunlight throughout the day. The molluscs were submerged
on July 7, 1947, and were examined for settled larvae at intervals thereafter.

During the summer of 1948 small wire bags (18 in. x 12 in.; see Pl. 1, fig. 6)
were filled with shells and rocks and set out at weekly intervals from July 16
through September 2. Another such collector was set out on October 5. Each
collector contained twelve Crassostrea virginica shells, two Mercenaria mercenaria
shells, six pieces of white marble, and six pieces of phyllite, dark gray in color.
The latter two materials were included to determine whether the color of the
background influences settlement of larvae. The collectors were suspended from
the floating dock of the Milford Biological Laboratory twenty yards from the
shore at mean low tide level in a position where they were shaded from direct

SPECIES

Clathrina (Ascetta)
clathra (Schmidt)

Clathrina (A scetta)
coriacea (Montagu)

Grantia (Sycon) compressa

Fabricius
Leucosolenia (A scaltis)
botryoides(Ellis and

Solander)

Sycandra sp.

Sycon (Sycortis) ciliata

(Fabricius)

Sycon raphanus Schmidt

Ute (Sycon) glabra
Schmidt

Oscarella (Halisarca)
lobularis (Schmidt)

Oscarella lobularis
(Schmidt)

Plakortis nigra Lévi

LIFE HISTORY STUDIES 81

TABLE 24
BREEDING SEASONS OF SPONGES*

BREEDING PERIOD

LOCALITY | AUTHOR

CLASS CALCAREA

Larvae released in mid-
March

Larvae released in
October

Eggs or larvae observed
in August

Larvae released in July

Larvae released in
August

Larvae released in
August

Bay of Naples
Luc, Calvados,
France

Le Portel,
France

Luc, Calvados,
France
Luc, Calvados,

France

Luc, Calvados,
France

Bay of Naples

CLASS DEMOSPONGIAE

Order Homosclerophora

Larvae released in
October

Luc, Calvados,
France

Schmidtlein,
1878

Topsent, 1887
Topsent, 1895

Topsent, 1887

Topsent, 1887

Topsent, 1887

Larvae released from Schmidtlein,
Dec. to March 1878
Larvae released in early | Bay of Naples Schmidtlein,
March 1878

Topsent, 1887

Topsent, 1895

Eggs or larvae observed Le Portel,
in August France
Eggs and sperm present | Red Sea Lévi, 1953b

in January.
Hermaphroditic

* Names in parentheses are those used in the reference cited.

(Continued)

82

MARINE SPONGES

TABLE 24—Continued

BREEDING SEASONS OF SPONGES*

SPECIES

BREEDING PERIOD

LOCALITY | AUTHOR

Order Epipolasida

Tethya aurantium

(Pallas) (T. lyncurium)

Tethya aurantium

(Pallas) (T. lyncurium)

Tethya aurantium
(Pallas)

Polymastia mammiullaris

(Miiller)

Polymastia robusta
Bowerbank

Larvae released at end of

May

Larvae released in
September

Eggs released in early
August

Bay of Naples

Luc, Calvados,
France

Roscoff, France

Order Hadromerina

Larvae released in
October

Luc, Calvados,
France

Schmidtlein,
1878

Topsent, 1887

Lévi, 1956

Topsent, 1887

Larvae released in
September

Suberites (Ficulina)
jicus (Johnston)

Suberites ficus

(Johnston)

Luc, Calvados,
France

Early autumn (beginning

in September)

Channel Coast
of France

Eggs released in early
October

Order Axinellida

Raspailia (Dictyocy-
lindrus) ramosa
(Montagu)

Eggs observed in October

Roscoff, France

Topsent, 1887

Topsent, 1900

Lévi, 1956

Luc, Calvados,
France

Sigmadocia (Gellius)
coucht (Bowerbank)

Tylodesma annexa

(Schmidt)

Vibulinus (Dictyocy-
lindrus) stuposus
(Montagu)

Topsent, 1887

Topsent, 1895

Eggs observed in August | Le Portel,

France
Eggs and sperm observed | Roscoff, Lévi, 1956
in September France

Eggs observed in
September

Luc, Calvados,
France

Topsent, 1887

(Continued)

SPECIES

Halichondria glabra
Bowerbank

Halichondria
(Amorphina) sp.

Halichondria sp.

Hymentacidon
(Amorphina) caruncula
(Bowerbank)

Hymentiacidon caruncula
(Bowerbank)

Hymentacidon sanguinea

(Grant)

Anchinoe? (Pronax)
plumosa (Montagu)

Carmia? (Esperia)
lorenzi (Schmidt)

Carmia? (Desmacidon)
similaris (Bowerbank)

Ligrota? (Microciona)
spinarcus (Carter and

Hope)

Microciona armata
Bowerbank

Mycale (Esperia)
lingua (Bowerbank)

Mycale? (Raphiodesma)

sordida (Bowerbank)

LIFE HISTORY STUDIES

TABLE 24—Continued
BREEDING SEASONS OF SPONGES*

BREEDING PERIOD

LOCALITY | AUTHOR

Order Halichondrina

Eggs or larvae observed
in August

Larvae released in
January

Larvae released in June
Larvae released in
September

August and September

Larvae released in
August

Roscoff, France

Le Portel,
France

Bay of Naples

Topsent, 1895

Schmidtlein,
1878

Luc, Calvados,
France

Channel Coast

of France

Roscoff,
France

Order Poecilosclerina

Larvae released in July,
August

Larvae released in
October, November

Larvae released in July

Eggs or larvae observed
in August

Larvae released in
September

Larvae released in
December

Larvae released in
September

Isle Verte,

Brittany, France

Bay of Naples

Lévi, 1956

Topsent, 1887

Topsent, 1911

Lévi, 1956

Lévi, 1956

Maas, 1892

Luc, Calvados,
France

Topsent, 1887

Le Portel,
France

Luc, Calvados,
France
Bay of Naples

Luc, Calvados,
France

Topsent, 1895

Topsent, 1887

Maas, 1892

Topsent, 1887

(Continued)

84

SPECIES

Mycale syrinx
(Schmidt)

Mycale (Esperia) sp.
Mycalecarmia?
(Esperella) littoralis

(Topsent)

Myxilla (Dendoryx)
incrustans (Bowerbank)

Myxtlla (Dendoryx)
incrustans (Bowerbank)

Myxilla radiata Topsent

Plumohalichondria
plumosa Arnesen

Stylopus cortaceus
(Fristedt)

Stylopus? (Leptosia)

dujardint (Bowerbank)

Tedantone foetida
Wilson

Acervochalina (Chalina)
gracilenta (Bowerbank)

Adocia? sp.
Desmacidon fruticosum
(Montagu)

Gellius angulatus
(Bowerbank)

MARINE SPONGES

TABLE 24—Continued
BREEDING SEASONS OF SPONGES*

BREEDING PERIOD LOCALITY

Larvae observed in
October

Larvae released in
December and January

Eggs or larvae observed

in August
Larvae released in
September

Eggs or lavae observed
in August

Eggs or larvae observed
in August

Eggs or larvae released
in August

Larvae released in
mid-September

Eggs or larvae released
in August

Eggs observed in
September and October

Bay of Naples

Bay of Naples

Le Portel,
France

Luc, Calvados,
France

Le Portel,
France

Le Portel,
France

Le Portel,
France

Isle Verte,

Brittany, France

Le Portel,
France

Bahamas

Order Haplosclerina

Larvae released in
September

Larvae released in
December and January

Larvae released in April

Larvae released in
September

Luc, Calvados,
France

Luc, Calvados,
France

Roscoff, France

Brest, France

| AUTHOR

Wilson, 1935
Schmidtlein,
1878
Topsent, 1895
Topsent, 1887
Topsent, 1895
Topsent, 1895
Topsent, 1895
Lévi, 1956

Topsent, 1895

Wilson, 1894

Topsent, 1887
Topsent, 1887
Lévi, 1956

Lévi, 1956

(Continued)

SPECIES

Haliclona indistincta
(Bowerbank)

Haliclona (Chalina)
oculata (Pallas)

Haliclona (Chalina)
oculata (Pallas)

Haliclona (Chalina)
oculata (Pallas)

Haliclona (Reniera)
rosea (Bowerbank)

Haliclona (Reniera)
simulans (Bowerbank)

Aplysilla sulfurea
Schulze

Halisarca dujardini
Johnston

Halisarca metschnikovi
Lévi

Ircinia (Hircinia)
acuta (Hyatt)

LIFE HISTORY STUDIES

TABLE 24—Concluded
BREEDING SEASONS OF SPONGES#*

BREEDING PERIOD

Larvae released from
May to July

Larvae observed from
June 15-22

Larvae released in July

Eggs or larvae observed
in August
Larvae released in July

Larvae released in
August

Isle Verte,

Brittany, France

Calvados,
France

Luc, Calvados,
France

Le Portel,
France

Luc, Calvados,
France

Luc, Calvados,
France

Order Dendroceratida

Eggs or larvae observed
in August

Le Portel,
France

Larvae released from
June to September

Larvae released about
June 1st

Luc, Calvados;
Roscoff, France

Roscoff, France

Order Dictyoceratida

Eggs observed in
September

Phyllospongia foliascens
(Pallas)

Spongia officinalis
Linné (Hippospongia
equina Schmidt)

Larvae released in
December and January

Bahamas

Red Sea

Larvae released from end | Tunis

of March to third week in

June; most common in
late May

* Names in parentheses are those used in the reference cited.

85

| LOCALITY AUTHOR

Lévi, 1956

Topsent, 1911
Topsent, 1887
Topsent, 1895
Topsent, 1887

Topsent, 1887

Topsent, 1895

Topsent, 1887;

Lévi, 1956

Lévi, 1956

Wilson, 1894

Lévi, 1956

Vaney and
Allemand-
Martin, 1918

86 MARINE SPONGES

TABLE 25
CULTCH PROVIDED FOR SPONGE LARVAE

NO. OF LIVING NO. OF DEAD
SPECIES OF MOLLUSC SPECIMENS SHELLS
SET OUT SET OUT
Crassostrea virginica (1 yr. old) 10 set-bearing None
shells
Crassostrea virginica (2 yrs. old) bei: 14 4
Crassostrea virginica (3 yrs. old) " 18 18
Crassostrea virginica (4 yrs. old) Ea 14 7
Crassostrea virginica (5 yrs. old) 2 3
Crassostrea virginica (totals) 58 32
Mercenaria mercenaria 4 12
Mya arenaria 12 12
Mytilus edulis 14 None
Spisula solidissima None 6
Arctica islandica None 12
Pecten irradians None 6
Ensis directus None 8
Crepidula fornicata None 12 ss
TOTAL 88 100

sunlight during the greater part of the day, with the exception of a short time in
the late afternoon. Examinations of the cultch were made twice monthly from
August 2 through December 11.

A study was also made of the depth of maximum settling of sponge larvae in
the harbor. For this work a wire cultch bag (PI. 1, fig. 6) was suspended from the
Milford Laboratory dock at each of the following depths: three feet above mean
low water, at mean low water, 214 feet below mean low water, and five feet below
mean low water (bottom of harbor). Each bag was provided with 12 Crassostrea
virginica shells and six Mercenaria mercenaria shells. The bags were submerged
on July 14, 1947, in a location which was shaded during most of the day.

Haliclona loosanoffi was the only sponge which settled on the cultch at this
site. On September 10, six shells bearing well-established colonies of this sponge
were removed from the bags and were attached to tiles supported horizontally in

LIFE HISTORY STUDIES 87

wooden frames so that the course of development of individual colonies could be
followed. One such frame bearing three shells with a total of 22 colonies attached
was suspended from the dock; another, bearing three shells with 17 attached
colonies was hung from the float. A record of growth and survival in these colonies
was kept, with further examinations on October 10 and December 5.

RESULTS AND DISCUSSION

Tables 26 and 27 list the total number of sponges of each species observed
at each examination.

TABLE 26
SETTLING OF SPONGE LARVAE, MILFORD HARBOR. SUMMER, 1947

DATE OF OBSERVATION
SPECIES OF SPONGE S| | Ss —_ cme

7/25 | 8/14 | 8/20 | 8/25 | 9/4 | 10/8 | 10/22

Halichondria bowerbanki Burton 0 0 — 3 -— — 127

Haliclona loosanoffi Hartman (on float) 0 0 — 0 — pet 29

Haliclona loosanoffi Hartman (on dock) 0 ~- 3 = 29 82 os

Microciona prolifera (E. and S.) cay 0 1 yoaine Rani, ae ae eae.

Cliona celata Grant 0 y — 2 ee ea 8

Cliona vastifica Hancock 0 0 elias aia peor nay apc
TABLE 27

SETTLING OF SPONGE LARVAE, MILFORD HARBOR. SUMMER, 1948

DATE OF OBSERVATION
SPECIES OF SPONGE > [| | | | |

Halichondria bowerbanki 0 a 10 10 9 12 12 <)

Burton
Haliclona loosanoffi 0 0 2 25 43 13 3 0
Hartman

Each species is considered individually in the following discussion.

Cliona celata Grant

Larvae of this species began to settle on calcareous substrata in Milford Har-
bor in early August and continued to do so through early October (Table 26).

88 MARINE SPONGES

Old (1942) found eggs in the clionids of Chesapeake Bay beginning about August
Ist; larvae settled on shells or calcite as early as July 28. Regrettably Old failed to
record the species of clionid on which these observations were made.

Table 28 shows the distribution of the eight colonies of Cliona celata which
settled on the cultch in the summer of 1947.

TABLE 28
SETTLING OF CLIONA CELATA LARVAE

SPECIES OF MOLLUSC AGE OF MOLLUSC NO. OF Cliona COLONIES
Mercenaria (alive) Sys. 1
Crassostrea (alive) 3 yrs. 2
Crassostrea (alive) 4 yrs. 1
Crassostrea (alive) aide 5 yrs. 3
Crassostrea (shell) 2yns: 1

Although the number of larvae which settled is unfortunately small, an appli-
cation of the chi-square test of independence to the data reveals that the associa-
tion of Cliona larvae with oysters is significant at the 2 per cent level. In this test,
infected (7 in number) and non-infected (83) oyster shells were compared with
infected (1) and non-infected (97) shells of other species. The tendency for settling
Cliona larvae to avoid oysters of the first two year classes was noted also by
Topsent (1900). Doubtless the thinness of the shells of young oysters is less favor-
able to growth of the sponge colony within them. It is noteworthy that clionids
are also rare in the thin shells of Ensis, Mytilus, and Volsella. It should also be
pointed out that in the present investigations the surface area of the 24 young,
living oysters was only about one-quarter of the surface area offered by the 34
larger living ones.

Of possible importance in explaining the apparent preferred settlement of
Cliona larvae on oysters is the fact that these shells are invariably provided with
corrugations and ridges which, together with the overhanging areas representing
regions of active growth in previous seasons, provide sheltered locations in which
the larvae can find protection from water currents during the critical periods of
metamorphosis and establishment of a burrow in the shell. The question of why
living molluscs attract the larvae is more difficult, though this tendency, if verified,
may be associated with the water currents created by living molluscs.

On the oyster beds in the New Haven region of Long Island Sound, the inci-
dence of infection of Crassostrea virginica (both living and dead) with boring
sponges of the genus Cliona is of the order of 80 to 90 per cent. Other shells con-
tributing to the cultch or occurring naturally on the beds and commonly bearing

LIFE HISTORY STUDIES 89

the sponge are Mercenaria mercenaria, Mya arenaria, Pecten irradians, Crepidula
fornicata, Busycon canaliculatum, and Busycon carica. Instances of the occurrence
of Cliona in shells of Ensis, Mytilus, and Volsella are rare.

Is the crystalline structure of the calcareous substratum of importance in ex-
cluding Cliona from some shells? A consideration of what is known about the
method of excavation is essential to a discussion of this question. Nasonov (1883)
made a very careful study of the excavating process in Cliona stationis. By collect-
ing the larvae on thin calcareous plates, he was able to follow the boring process
from its inception. A day or two after settling, the sponge colony had grown into
a round, flat plate about 0.7 mm. in diameter, and at this time the excavating
activities began. Processes of sponge tissue were sent down into the substratum
dissolving the calcareous matter before them and forming initially a rosette-like
pattern. At the end of the second day it was found that small hemispherical frag-
ments of the calcareous matter were extruded from the sponge, and the colony
had moved into the depression so formed. Since no spicules were present in the
colony at this stage of development, Nasonov gave the first conclusive evidence
against the mechanical theory of Hancock (1849, 1867). In his earlier paper this
author described the existence of siliceous bodies, about 42u across, imbedded in
the surface of Cliona and suggested that “every portion of it will cut with the
keenness of glass-paper.” Later (1867) he reported that the supposed siliceous
bodies were in reality fragments of conchiolin from the oyster shell being exca-
vated. He then suggested that smaller angulated siliceous bodies (about 4u across),
also imbedded in the surface, or the spicules themselves, “must’’ accomplish the
boring process. It is possible, of course, that both chemical processes and mechani-
cal pressure brought about by the contractility of the sponge enter into the forma-
tion of burrows.

Old (1942) confirmed Nasonov’s observations on the boring activities of re-
cently settled clionid larvae, pointing out that many of the young sponges lack
spicules when they first begin to perforate shells. He found that the sponge is
able to penetrate the thin layers of conchiolin occurring between the layers of
calcium carbonate in the shell. The conchiolin is removed in tiny pieces corre-
sponding in shape to the expelled shell fragments. Although he suggests that this
observation argues against the obvious hypothesis that an acid secretion is in-
volved in the boring process, he also reports a slight daily increase in dissolved
calcium in sea water containing actively boring clionids. Nasonov (1924) noted
that newly settled larvae of Cliona stationis can perforate the periostracum of
oysters in the same manner as they attack the calcareous portions of the shell. He
found that the larvae also penetrate the chitin associated with the calcareous
plates of Balanus. Nasonov suggested that the sponges probably produce an en-
zyme which dissolves conchiolin as well as an acid secretion which erodes calcium
carbonate.

The hemispherical fragments of calcareous matter which Nasonov noted have
been observed subsequently by many investigators. These fragments can be seen
issuing from the oscules of Cliona celata; and when infected oysters are kept in a
quiet aquarium for several days, piles of such fragments accumulate at the base of

1 Nikitin (1934) reports that Cliona stationis is found only in the shells of Ostrea on the
Gudaut Oyster Bank of the Black Sea coast of the Caucasus near Sukhumi. He goes on to
say that “in the other regions of the Black Sea Cliona is met with sometimes also upon
Mytilus.”

90 MARINE SPONGES

each oscular tube.2 The surface of the internal cavities of an infected shell is
pitted, indicating that numerous such fragments have been removed in the ex-
cavating process. These pits vary from circular to elliptical in basal cross-section,
and their configuration in the shells of both Crassostrea and Mercenaria is similar
to that observed by Nasonov in calcite. The shapes and dimensions of the basal
cross-sections in a Crassostrea virginica shell were found to vary from an ellipse
with axes of 45u and 32u to an ellipse with axes of 70u and 58y; circles of 45 to
50u in diameter occurred most frequently. In a shell of Mercenaria mercenaria the
shapes and dimensions varied from a circle 25u in diameter to an ellipse with axes
of 82u and 50u; circles with diameters of 45u to 50u again predominated. The
general similarity of the boring process in the cases cited make it seem unlikely
that the detailed structure of the calcareous substratum greatly influences the
mechanism.

A curious observation by Cotte (1914) is suggestive of an exclusion of Cliona
based on the composition of the substrate. He has reported a relationship between
Cliona viridis and Lithophyllum expansum, in which the sponge is incapable of
excavating the calcareous skeleton of the alga in order to send the characteristic
tubules to the surface of the substratum. Instead, long, imperforate tubes of the
coralline alga form; these are filled with sponge tissue and provided with a single
apical sponge tubule through which excurrent and incurrent streams of water are
maintained. The enriched supply of carbonic acid in the vicinity of the terminal
oscule probably assists the alga in laying down more of the calcareous covering.
It is possible that the dolomitic structure of the alga excludes Cliona in this case.
The calcareous skeleton of coralline algae is admixed with the maximum amount
of MgCO; known in any organisms [up to 36 per cent of the ash in some species
(see Vinogradov, 1953)]. Novatéek (1930) found 6.27 per cent MgO (=13.11%
MgCoOs) in the ash of Lithophyllum expansum.

Vosmaer (1933) noted that large portions of colonies of Lithophyllum in-
fected by Cliona celata are sometimes imperforate, but he also described cases in
which the alga had been completely destroyed by the sponge. Volz (1939) observed
nodules of coralline algae at Rovigno which were completely filled with sponge
tissue and found that tubules of the sponge readily penetrate the outer covering
of calcareous matter. He noted also that stages exist in which the calcareous mate-
rial has been dissolved completely, leaving free-living colonies of the sponge.
Similar associations of Cliona celata with coralline algae (Lithothamnion sp.) are
common on the coast of central California in the experience of the present author;
in no case, however, are colonies found in which tubule formation is inhibited as
described by Cotte. Clionids, therefore, are certainly capable of excavating the
dolomitic skeletons of corallines; the factors involved in Cotte’s example remain
obscure.

Topsent (1900) has noted that clionids do not infect the Portuguese oyster
(Ostrea angulata), which has become established in places along the coast of
France. It is of interest to note in this connection that Arndt (1941) failed to find
any clionids on the coast of Portugal.

As the growth of Cliona celata colonies continues, the sponge eventually over-
grows the shell and still later the shell disappears entirely, leaving a free-living

2 This observation provides a clue as to whether the tubules of Cliona are functionally
specialized. The restriction of such piles of calcareous fragments to the bases of tubules
bearing a single terminal osculum provides evidence against the supposition sometimes
held that tubules of the second type have both incurrent and excurrent functions.

LIFE HISTORY STUDIES 9]

sponge colony which increases in size indefinitely if undisturbed. One such colony,
dredged on an oyster bed off Momauguin Beach, East Haven, Connecticut, where
the bottom had been undisturbed for a period of ten years, had reached the ex-
traordinary dimensions of 35 cm. x 16 cm. x 8 cm. Such specimens were formerly
included in different genera (Papillina suberea Schmidt or Raphyrus griffithsii
Parfitt).

An experiment was set up during the course of the present investigations to
determine whether or not Cliona celata can reverse its life history. Cubes of tissue
of a gamma-stage sponge (each about one cubic centimeter in volume) were at-
tached to uninfected oyster shells. Similarly pieces of oyster shell infected with
the alpha-stage of Cliona celata (each shell fragment having a volume of about
2 cm.) were attached to uninfected oyster shells. All were placed in a wire basket
and suspended on August 22, 1947, from the dock of the Milford Laboratory at a
depth midway between the bottom and mean low water (depth: 6 feet at m.l.w.).
Examinations of the growth of the colonies were made at intervals thereafter.
After one month of growth there was no evidence of penetration of the new
shells in any case. ‘The gamma-stage fragments had regenerated their cut surfaces
and were now firmly attached to the shells on which they had been planted. Later
examinations were made on October 27 and on December 17; the results at these
times are shown in Tables 29 and 30.

In the gamma-stage, Cliona celata retains the capacity of boring into calcareous
matter with which it comes into contact. It is apparent, however, that the penetra-
tion of the new substratum was accomplished more quickly by sponges of the
alpha-stage than by those of the gamma-stage. This is unexpected especially in

TABLE 29

PROGRESS OF GROWTH IN THE ALPHA-STAGE TRANSPLANTS

SIDE OF SHELL PENETRATION INTO NEW SHELL*

SHELE AGE OF | oN WHICH SPONGE
Ne: SHELL WAS PLANTED

October 27 December 17
1 2 yrs. Lower 12—upper surface 20-upper surface
2 2 yrs. Upper 1—lower surface 3-lower surface
3 3 yrs. Lower 9-upper surface 12-upper surface
4 Suyls: Upper 13-lower surface 24—lower surface
5 4 yrs. Lower 7—upper surface 22—upper surface
6 4 yrs. Upper Sponge fragment lost —
al 5 yrs. Lower 18—upper surface 18—upper surface
8 6 yrs. Upper None-lower surface 3-lower surface

* In terms of number of holes in surface of shell on side opposite that on which sponge was planted.

92 MARINE SPONGES

TABLE 30
PROGRESS OF GROWTH IN THE GAMMA-STAGE TRANSPLANTS

SIDE OF SHELL PENETRATION INTO NEW SHELL*
eee Bo oy WwHICHisEo Nee |
ae SHELL WAS PLANTED
October 27 December 17

1 2 yrs Upper Sponge fragment lost —

; 3 yrs. Lower Sponge fragment lost =

3 3 yrs. Upper No evidence of 3-lower surface
penetration

4 4 yrs. Lower No evidence of 6—-upper surface
penetration

5 4 yrs. Lower No evidence of None-upper surface
penetration Many-under sponge,

lower surface

6 5 yrs. Upper No evidence of 4—lower surface
penetration

ff 5 yrs. Lower No evidence of None-upper surface
penetration Many-under sponge,

lower surface

8 6 yrs. Upper 2-under sponge, 6-lower surface

upper surface

* In terms of number of holes in surface of shell on side opposite that on which sponge was planted.

view of the fact that more sponge surface is in contact with the shell in the case
of the gamma-stage transplants, and one would suppose that immediate penetra-
tion of the shell would ensue. Instead, as Table 30 indicates, there were no cases
of excavation through the entire thickness of the shell by the gamma-stage frag-
ments even two months after affixation, whereas in every case but one, the alpha-
stage fragments had accomplished a complete penetration of the shell so that from
one to 18 holes were present on the side of the shell opposite that on which the
transplant had been attached. On December 17, four of the six gamma-stage
transplants had bored completely through the shells with a mean of five holes
(range, three-six) whereas all seven of the alpha-stage transplants had penetrated
completely with a mean of 15 holes (range, three—24). Why there should be a
longer latent period in the case of the gamma-stage colonies is difficult to say;
perhaps the delay simply reflects a period necessary for the regeneration of the
cut surfaces of the free-living samples.

Although the gamma-stage will grow either as a free-living colony or revert to
the boring habit, it has, to the writer’s knowledge, never been demonstrated that
larvae can grow directly into the gamma-stage without passing through a boring or

LIFE HISTORY STUDIES 93

alpha-stage. Seemingly in the early stages of growth, the colony requires the pro-
tection provided by mollusc shells and other calcareous matter into which it bores.
It would be instructive to attempt to rear larvae under laboratory conditions in
the absence of calcareous substrata to find out whether attachment takes place on
other substances and how far development proceeds under these conditions. On
the other hand, small portions of colonies already established in oyster shells might
be isolated from their calcareous substratum to find out if survival in the free-
living stage is dependent upon size or age of colony. During the summer of 1949,
small cubes (one cm.*) of gamma-stage tissue were planted on phyllite slabs; their
growth was followed for three months after transplantation. Growth was slow
and consisted mainly of a spreading out over the rock from the base of the
fragment.

In Vosmaer’s exhaustive account of the genus Spirastrella (1911) he mentions
certain forms of the highly variable species,? S. purpurea, which dwell in cavities
in calcareous matter. There is no evidence in these instances that the sponge col-
onies have excavated the cavities; they simply inhabit abandoned worm tubes and
other similar situations. These forms are so characteristically associated with
cavities that many authors have mistaken them for true boring sponges. Other
forms of what Vosmaer regarded as the same species live on the surface of corals
and shells, lacking the tendency to dwell in cavities. This observation provides a
clue to the origin of the boring habit. It is not unlikely that a cavity-dwelling
tendency resulted in the establishment of a process by which the cavities could
be enlarged. After this process had appeared, the sponge was capable of estab-
lishing its own cavities and was no longer restricted by fortuitous settling in
naturally occurring ones.

Cliona vastifica Hancock

Only a single colony of Cliona vastifica settled on the shells set out. It occurred
in the shell of a five-year oyster and had probably settled some time during the
month of September, the first date of observation having been October 13, 1947,
at which time the colony was well established. Topsent (1900) found individuals
of this species filled with orange unicellular eggs in late September in the English
Channel. Reproduction would therefore seem to be somewhat later there. In the
New Haven Harbor region of Long Island Sound this species is never as abundant
as Cliona celata, a fact borne out in the Milford Harbor larval settling observa-
tions.

Cliona vastifica has never been observed to overgrow its shell and assume a free-
living condition. This is surprising since a point must be reached in the develop-
ment of the sponge colony when the shell in which it is growing is completely
occupied by sponge tissue, so that further growth can result in but one thing:
overgrowth of the shell. It is impossible to say at present whether this observed
difference between the two species, Cliona celata and C. vastifica, is a result of a
slower rate of growth or perhaps a shorter life span in the latter form.

An interesting mode of asexual reproduction was observed in specimens of
Cliona vastifica which had been kept in aquaria in the Milford Laboratory for
several weeks during the summer of 1947. Protruding from certain of the holes in
the shell harboring the sponge were long, thin tubes bearing bulbous expansions
along their length. It is uncertain whether these tubules possessed any incurrent

8In a later monograph, Vosmaer (1933, p. 334) was inclined to regard Cliona and
Spirastrella as synonyms.

94 MARINE SPONGES

or excurrent function, but it seems most likely that the rounded bodies were occa-
sionally freed from their attachment and thus served to establish new colonies in
other shells. Indeed, it is possible that these structures are provided with cilia at
the time of their release much as the asexually produced larvae described by
Wilson (1894). This method of reproduction was observed by Merejkowsky (1879)
in Rinalda arctica (—Polymastia mammillaris) occurring in the White Sea and
was subsequently observed by Arnesen (1917) in the same species. It is probably
not an uncommon method of reproduction in tubule-bearing species of Hadro-
merina, but it has been very generally overlooked.

Perhaps the most interesting problem presented by Cliona vastifica is its oc-
currence along with the commoner Cliona celata and the rarer Cliona lobata in
what seems to be a competitive relationship. A further consideration of this matter
will be deferred until the results of salinity tolerance experiments in the several
species have been discussed.

Haliclona loosanoffi Hartman

This sponge is one of the most common species found in Milford Harbor, and
comparatively large numbers of settled larvae were observed during both seasons
of study, providing an opportunity to obtain information on the period of larval
settling as well as the survival of the young colonies. In 1947 data regarding this
species were secured from two sources: (1) the molluscs in the tray suspended from
the float in Milford Harbor, (2) the cultch bags set out at varying depths from the
dock.

Haliclona loosanoffi larvae began settling in mid-August in 1947 and in mid-
September in 1948. In both years the peak of settling had been reached by early
October as is shown in figure 28.

The growth of 39 colonies of Haliclona loosanoffi was followed in greater de-
tail from September 10, 1947 to December 5, 1947. These observations are listed
in Table 32 and summarized in Table 31.

An attempt has been made to construct a life table for Haliclona loosanoffi
although the data are meager. It must be emphasized, however, that this analysis
pertains to only one part of the life cycle, extending from the time of establish-

PABLE *31
SURVIVAL OF HALICLONA LOOSANOFFI

NO. OF NO. OF NO. OF
LOCATION NO. OF NO. OF COLONIES | COLONIES | COLONIES NO. OF
OF SPONGES COLONIES | COLONIES WHICH WITH WITH |GEMMULES
SETTLED | GEMMULES | GEMMULES
SINCE

Sept. 10 Oct. 10 Sept. 10 Oct. 10 Dec. 5 Dec. 5

Sponges on dock 21 9 1 1 6 440

Sponges on float 18 10 1 1 6 105

TOTAL 39 19 2 2 12 545

LIFE HISTORY STUDIES 95

ment of the young sessile colonies which have recently metamorphosed from free-
swimming larvae up to the time of gemmule formation and winter killing. A
source of error results from the fact that the exact date of settling is unknown for
the 39 colonies studied. It can be stated only that they settled some time beween
August 20 and September 10, a three-week period which may represent the spread
in their ages. Survival was recorded on September 10, October 10, and December 5.
On November 10 it was observed that all colonies had died back, but no detailed
record of gemmulation was kept. Presumably the observations on that date would
have been identical with those on December 5, although fragmentation of the
gemmules following death of the colony is not impossible.

TABLE 32
DATA ON GROWTH OF SELECTED COLONIES OF HALICLONA LOOSANOFFI

COLONY VOLUME VOLUME RATIO OF NO. OF
SHELL NO. NO. IN. DEX} INDEX VOLUMES GEMMULES
Sept. 10 Oct. 10 Dec. 5
BAe Pac hse teiiay He pals 1 570 3850** 6.8 350
(831)* Ge S)s

2 -— 84 —. a

18s. ke ee 1 319 — — a
2 158 — a —=

3 94 _ — ==

4 27 108 4.0 20

5 27 — — —

6 ue _ — —

if 87 _ — =

8 27 = — ==

9 27 — _ —

10 Dil -- — —

11 54 a -— —

12 27 i — —

13 32 173 5.4 —

POA as bas fi 248 450** 1.8 _

(241)* Gn0)4

2 8 os —- —

3 98 54 0.5 ile |

4 180 — -- —

5 215 206 0.8 37

6 106 958 9.0 14

wh 196 178 0.9 8

IS | Ae eee if 36 a _. _-
2 4 — — _

3 51 140 pat 20

4 198 248** 15 —

5 36 32 0.9 —

(Continued)

96 MARINE SPONGES
TABLE 32—Concluded

COLONY VOLUME VOLUME RATIO OF NO. OF
SHELL NO. NO. INDEX T INDEX VOLUMES GEMMULES
Sept. 10 Oct. 10 Dec. 5
II-A (Cont.) 6 105 —

| 27 60 22 9
8 24 — — —
9 18 42 2-3 —
10 93 — — —
11 —= 48 = 7
PB els ts eee oe 1 3 — = —
2 75) — == —
3 150 Dili Oz 23
4 193 3875 20.0 —
5 88 — aa =
|) Ee Oa Mehdi pes caccne i 1 342 288 0.8 —
2 27 421 15.6 32
3 24 25 1 ot 14

* Living tissue. Rest of colony dead.
** Gemules present on Oct. 10.
{ Volume index = product of length X width X height. Mean ratio of volumes — 4.2
I—Shells so marked were suspended from dock.
II—Shells so marked were suspended from float.

In spite of the weaknesses of the data a life table is presented (Table 33) in
order that survival may be compared with data existent for other sessile marine
organisms (Deevey, 1947).

TABLE 33
LIFE TABLE FOR NEWLY SETTLED COLONIES OF HALICLONA
LOOSA NOFFI
x ih 1 1000 gq:
NO. DYING IN NO. SURVIVING AT MORTALITY RATE
AGE IN MONTHS INTERVAL OUT OF BEGINNING OF AGE PER 1000 ALIVE
1000 SETTLED INTERVAL OUT OF AT BEGINNING OF
LARVAE 1000 SETTLED LARVAE AGE INTERVAL
0-1 564 1000 564
1-2 154 436 353

2-3 0 282 0

LIFE HISTORY STUDIES 97

Data on the number of free-swimming larvae which are produced and the per-
centage of these which settle successfully will be necessary to complete the picture,
but such are unavailable at present. As it stands, the survivorship curve approaches
that of Pearl and Miner’s (1935) positively skew rectangular type in which there
is heavy mortality in early life with long survival of the few individuals which
live to advanced ages.

Probable causes of death of newly settled sponges are several and are de-
pendent in part upon time. Predation by gastropods is very probably an impor-
tant cause of mortality. Nudibranchs have long been suspected of browsing on
sponge tissue, and recently Burton (1949) has demonstrated that snails and
limpets do not pass by this spiculous fare. He observed spicules in the guts of
Littorina littorea and L. obtusata as well as Patella, and found grazed areas on the
sponge colonies from which the molluscs were removed. The present writer has
observed that chitons likewise feed on sponges on the central California coast.
Crowding by faster growing organisms may be of some importance, but on the
whole, sponges are able to hold their own by encrusting such organisms as tuni-
cates, bryozoans, and barnacles. In certain cases coalescence of several colonies
may occur and introduce an error. And in other instances parts of a large colony
will degenerate leaving several isolated portions to continue; these cases are easily
detected, however, by the continuity of the skeletal framework. Some of the
sponges which survive until late October without having produced gemmules,
will succumb to decreasing temperature conditions, a problem discussed at greater
length in a later section.

Data on the winter survival of gemmules are also unavailable. The gemmules
reported here were kept in a tray in Milford Harbor during the winter of 1947-48
in anticipation of securing information on their survival. However, when the
Shells to which they had been attached were examined in the spring, no trace of
the gemmules could be found. Presumably they had all sloughed off during the
rigorous winter of that year. Provided that the gemmules remain attached to their
original substratum during the entire winter, it is likely that the same number of
colonies would start growth in the spring as had formed gemmules in the fall. The
germination of but one gemmule would suffice to renew the growth of the colony,
and the germination of many gemmules (from the same fall colony) would lead to
a single, large colony since the young sponges produced from the individual gem-
mules would doubtless coalesce. In such a case, the survivorship curve could be
continued as a straight line up to the time of early growth of the sponge colonies
the following year.

If, on the other hand, the gemmules regularly slough off, as was the case with
those under observation in Milford Harbor, then renewed growth in the spring
must depend upon the number of gemmules which come to rest on a suitable
substratum at the time of germination. A great increase in the number of colonies
could result from this series of events, as is shown by the 1947 data where 12
colonies produced 545 gemmules. Another uncertainty which arises in regard to
the role played by gemmules in increasing the population in this species, concerns
the type of organism produced upon germination. Observations of gemmules
which germinated during the winter in the laboratory revealed that sessile colonies
arise directly from the gemmules in this case; however, it has been shown by
Wilson (1894) that the gemmules of some monaxonid sponges give rise to ciliated
larvae essentially like those produced sexually. If this is an alternative mode of
germination in Haliclona loosanoffi, the gemmules are unquestionably important

98 MARINE SPONGES

agents of population increase. A solution to these problems awaits more careful
and complete observations on the life history of this interesting sponge.
Reference to the graph in figure 28 indicates that the larvae settling on shells
suspended from the dock considerably outnumbered those settling on shells hung
from the float. Since the shells in the latter location outnumbered the former, so
that there was actually more surface area available for settling, it seems reasonable
to expect a correlation with some environmental factor. The only obvious differ-

80

60

NUMBERS OF COLONIES

40

rm)
a
2
ro)
a
°
°
w
ro}
2)
x
Ww
a
=
=)
2

Si

ILL

20

LEGEND
O——O Total no. of colonies settling on shells on dock

O--=-0 Total no. of colonies settling on shells on float.

LLLLLLLLE
CLMMMLAYA

ws

O-wO No. of colonies which produced gemmules.

28 25 4
July Aug. Sept. Oct. Dec.

Figure 28. Larval settling of Haliclona loosanoffi in two successive years. Milford
Harbor. Left, 1947; right, 1948.

@

22 10 N No. of colonies which settled since last date
XN of observation.

ence in the two locations concerns the amount of sunlight which reached them.
The float received the full light of the sun all day whereas the shell bags, sus-
pended from the dock, were shaded. In the other localities in Long Island Sound
where this species has been taken, the colonies are invariably located on the
undersides of overhanging granite blocks or on the lower sides of intertidal rocks
and boulders. ‘They have never been taken in exposed situations as is true of
Halichondria bowerbankt. The numbers of colonies of the two species, Haliclona
loosanoffi and Halichondria bowerbanki, which settled in exposed and shaded
situations in the two successive years are shown in Table 34.

TABLE 34
INFLUENCE OF LIGHT ON SETTLING OF SPONGE LARVAE

LARVAE SETTLING | LARVAE SETTLING | LARVAE SETTLING
SPECIES OF SPONGE IN FULL SUN IN SHADE IN SHADE
1947 1947 1948

Halichondria bowerbanki

(Max. no. settling) 64 0 12

Haliclona loosanoffi
(Max. no. settling) 29 82 43

LIFE HISTORY STUDIES 99

On the other hand, an anlysis of settling on white marble as opposed to dark
gray phyllite fails to show any tendency for one to be favored over the other.
Comparison of settling on both of these substrata, as well as on shells of Crassostrea
virginica (fig. 29) reveals that settling on the several types of backgrounds was es-
sentially random. In the figure, half the total number of colonies which settled on
oyster shells is shaded so as to permit comparison with similar surface areas on
marble and phyllite.

It is of interest to note that McDougall (1943) found that Adocia tubifera was
the only organism which settled in his light box at random. McDougall pointed

OYSTER SHELLS MARBLE PHYLLITE
AN)

tion

(ae ey ee (te! So I9erets 5g) 2
Sept. Oct. Nov. Oct. Nov. Oct. Nov.

FicurE 29. Larval settling of Haliclona loosanoffi in relation to color of substra-
tum. See text.

NUMBERS OF COLONIES

out that the glass top on his box eliminated ultra-violet radiation which may be
the injurious element of sunlight for some organisms. However, the amount of
ultra-violet light which penetrates turbid coastal waters is probably insignifi-
cantly low, at least at a depth of 114 meters at which these organisms settled.
Consequently, it is likely that a basic difference in phototactic responses exists in
these two species.

The distribution of Haliclona loosanoffi colonies with depth is shown in
Table 35. Their exclusion from the shells on the bottom is doubtless a result of
the heavy silting in the Harbor so that only those colonies could survive which
settled on substrata suspended off the bottom where water currents were more
likely to be effective in preventing silt accumulation. The upper limit of settling
found here agrees with field observations elsewhere in the Sound where, in gen-
eral, mean low water represents the shoreward extreme of distribution.

Haliclona loosanoffi colonies were observed to die back in mid-November each
year from 1947 through 1949. The regularity with which the colonies perished
each fall suggests that an environmental factor is limiting their growth. The de-
generative process actually begins in late October or early November when the
living tissue of portions of the colonies dies back leaving the bare skeletal frame-
work. This degeneration proceeds slowly through the next few weeks until all
sponges have died by the end of the second week of November. In figure 30, the
surface water temperatures taken at high tide in Milford Harbor during the
months of October and November for the past three years are plotted along with
observations on the survival of colonies of Haliclona loosanoffi. In 1947 Haliclona
colonies were observed to be growing normally on October 10 and 22, but by

100 MARINE SPONGES

TABLE 35
LARVAL SETTLING OF HALICLONA LOOSANOFFI IN RELATION TO DEPTH

NUMBER OF COLONIES
DEPTH en es er Ore re

August 27, 1947 | September 4, 1947 | October 8, 1947

3 feet above mean low
water

Mean low water level — 4 37

2.5 feet below mean low

ae 3 24 45
5 feet below mean low
water — 1 —

November 10, all had died back leaving the skeleton and gemmules. During this
period the water temperatures had fallen from 17° C. to 11.6° C. In the following
year the colonies were thriving on October 5; by October 19 one colony had be-
gun to die back. On November 2 only four (out of 42) colonies were still in a

22
Q O---0 1947 im Open areas within figures
sie refer to living colonies.
Ai Oe i Pe ee Co-—oO | :
20 pes Make A Solid areas refer to
3 O------O 1949 @ dead colonies.

ww Q
@ z
5 (I) oar
2
=
z
ul
Oo 16
n
WJ
lJ
©
lw 14
ras)
i
ul
S12
te
<t
c
tu i
a
= 10
Ww
| ed

8

is) 10 15 20 25 30 4 9 14 19 24 29 4
October November

Figure 30. Graph comparing Milford Harbor water temperatures (surface, high
tide) with onset of degeneration in colonies of Haliclona loosanoffi over a three-
year period.

LIFE HISTORY STUDIES 101

healthy condition; five others were partially alive. All the rest had died back com-
pletely with only four of these leaving gemmules to carry them over the winter.
On November 18 all colonies were dead; nine had formed gemmules. In 1949 a
few colonies showed indications of degeneration on October 27; this process was
more extensive by November 2; all colonies were dead on November 16.

It is apparent from the graph that each year as the temperature fell below
16° C., degeneration began. It is impossible to set the limiting temperature very
exactly without experimental data; but the critical range certainly lies between
16° and 12°C.

Since this sponge is able to survive the winter in the gemmule state, winter
killing in itself is probably not a factor in limiting northward distribution. The
minimum temperature for reproduction would seem to be of greater importance
in this regard. The first date on which newly settled colonies of this sponge were
recorded during the three summers of observation is distinctly correlated with
the water temperatures of the season in question as can be seen in figure 31. Pre-
cise information about the timing of the early processes of the reproductive cycle
are unfortunately not available; nor are there any experimental data on the effect
of temperature on the reproductive cycle. It is therefore futile to postulate exact

26

24

22

20

TEMPERATURE — DEGREES CENTIGRADE

O---0 1947
10 | O—o 1948
Oreo 1949

Bold segments of graphs equal
periods of larval settling.

June July Aug. Sept. Oct. Nov. Dec.

Figure 31. Graph comparing Milford Harbor water temperatures (surface, high
tide) with onset of larval settling in Haliclona loosanoffi over a three-year period.

limiting temperatures in this discussion. However, the data at hand imply that
there is a certain minimum temperature (probably between 20-22° C.) at which
the reproductive process begins, and that the annual variation in appearance of
this temperature is reflected in the observations on larval settling.

102 MARINE SPONGES

Perhaps the length of growing season which remains after the larvae have
settled and metamorphosed is of equal importance in limiting the northward dis-
tribution of this sponge as the minimum temperature for repopulation. The
former time interval is, of course, dependent upon the latter; its importance con-
cerns the number of gemmules which are available for repopulation in the spring.
Table 36 indicates that the number of sponges which formed gemmules in 1947
was greater than that in 1948, and although these data are found to be insignifi-
cant when the chi-square test is applied, the number of gemmules formed per
sponge in 1947 was distinctly higher than that in 1948.

TABLE 36
GEMMULE FORMATION IN SUCCESSIVE YEARS

YEAR OF NO. OF COLONIES NO. OF COLONIES
OBSERVATION OBSERVED WITH GEMMULES
1947 39 12
1948 42 9

Exact counts of the gemmules produced in 1948 were not made, but the num-
ber was surely no higher than one-fourth of the 545 gemmules produced by 12
colonies in 1947. It seems likely that this difference resulted from the longer
growth period in 1947 with the consequent larger size of the colonies.

Halichondria bowerbanki Burton

Young Halichondria bowerbanki colonies were first observed during the last
week of August in 1947 and in early September in 1948. Settling continues from
that time through October and in 1948, at least, it extended into late November or
possibly early December, three colonies having settled between November 18 and
December 11 (see fig. 33). Subsequent survival of the colonies proved to depend
upon the severity of the winter. During the rigorous winter of 1947-48, all colonies
were killed back in Milford Harbor. On the other hand, during the mild weather
of the 1948-49 winter season, the colonies which had settled in the fall of 1948
survived and continued growth slowly. Some of these colonies were still growing
in the Harbor in mid-December of 1949, though all had noticeably declined in
vigor toward the end of their second fall season.

Essentially similar observations in regard to the importance of winter-killing
in determining the abundance of Halichondria bowerbanki in inshore locations,
were made at Momauguin (East Haven) and Pine Orchard (Branford), Connecti-
cut. In the former locality, large colonies of Halichondria bowerbanki were abun-
dant during the summer of 1947 on the reefs surrounding Stony Island offshore
from Momauguin Beach. During the following summer, however, such large col-
onies of this sponge were nowhere to be seen in this vicinity. There were numerous
small colonies encrusting the algae and rocks in the vicinity suggesting that the
area had been recolonized by asexually or sexually produced larvae from colonies
which had survived the severe winter of 1947—48 in deeper, offshore waters. During

LIFE HISTORY STUDIES 103

NUMBERS OF COLONIES

Be), See eee ep he) rE Ns! It
Aug. Sept. Oct. Nov. Dec.

NUMBERS OF COLONIES

O——O Total number of colonies.

No. of colonies which settled
since last date of observation.

WZ.

Ficure 33. Larval settling of Halichondria bowerbanki in two successive years.
Milford Harbor. Left, 1947; right, 1948.

104 MARINE SPONGES

the summer of 1949, conditions similar to those in 1947 prevailed with large sized
colonies abundantly in evidence. These colonies had doubtless lived throughout
the mild winter of 1948-49 and were ready to start rapid growth with the onset of
warm weather in the spring.

In the fall of 1948 a large number of young colonies of Halichondria bower-
banki were observed growing between extreme low water neaps and extreme
low water springs on ‘Tub Island, off Pine Orchard, Connecticut. These young
sponges measured from two to five centimeters in diameter and were grow-
ing at a density of about three per square meter on the northern end of the
island, frequently in exposed places on the bare granite surface. The fol-
lowing spring, virtually all these colonies had disappeared except for the few
which had settled in crevices, under overhanging rocks, or in other protected
spots. Winter killing and predation by snails probably account for the mortality
of the greater number of these colonies, but it is noteworthy that some of the
colonies were able to survive through the mild winter of 1948-49.

A discussion of the relation of light to settling in this species has already
been undertaken.

In regard to the intertidal distribution of this sponge, it is of regular occur-
rence in tidal pools and on the surfaces of rocks at Lighthouse Point (New Haven),
Double Beach (Branford), Pine Orchard (Branford), Stony Creek, and Hammon-
asset, Connecticut. It seldom survives higher than the level of extreme low tide
neaps, but in one instance, a thriving colony was observed in a tide pool at Double
Beach located just above the mid-tide level. With this one exception, these ob-
servations on the intertidal distribution of Halichondria bowerbanki agree with
those of Burton (1949) on H. panicea at Tor Bay, England.

Microciona prolifera (Ellis and Solander)

Microciona prolifera is an abundant sponge on the oyster beds of Long Island
Sound; it is surprising that so few were observed in the present investigations.
Reference to Table 26 indicates that it began settling in early August and con-
tinued up to mid-October. McDougall (1943) found essentially the same settling
period for this species in Beaufort Harbor, North Carolina. It is possible that
this species, like Halichondria bowerbanki, prefers well-lighted situations, for it
was not found on any of the cultch set out from the dock in a shaded location. It
is certainly true that intertidal and shallow water specimens of this sponge
commonly grow in exposed situations and are not restricted to the undersides of
rocks as is true of Haliclona loosanoffi.

SUMMARY

1. In this investigation Cliona celata larvae showed a significant tendency to
settle preferentially on shells of oysters. It is suggested that the many irregu-
larities of the surfaces of these shells favor settlement of clionid larvae.

2. The massive gamma-stage of Cliona celata is capable of reverting to the boring
habit when cuttings are applied to a calcareous substratum.

3. The formation of strands of spherical buds was observed as a mode of asexual
reproduction in Cliona vastifica.

4. A life table constructed from a limited amount of data on the survival and
death rates of newly settled Haliclona loosanoffi colonies shows that it is of
the positively skew rectangular type and is allied to that of other sessile
marine organisms such as barnacles.

10.

LIFE HISTORY STUDIES ' 105

. Larvae of Haliclona loosanoffi tend to respond negatively to light, contrary

to the larvae of Adocia tubifera, observed by McDougall at Beaufort, North
Carolina, which are unaffected by light in choosing a place to settle.

. The relationship of depth to larval settling was studied in Haliclona loosan-

offi. Heaviest settling occurred on shells at mean low tide level and on those
suspended freely in the water at 214 feet below mean low water. No settling
occurred at the bottom.

. Data accumulated over a period of three years reveal that in Haliclona

loosanoffi, the time of settling of larvae fluctuates annually in relation to
temperature; larvae are released shortly after the highest water temperatures
of the year.

. Degeneration of newly settled colonies of Haliclona loosanoffi also fluctuates

annually in relation to water temperature. In general, temperatures of 16° C.
or 15° C. lead to the inception of degeneration.

. It is suggested that two factors operate in limiting the northern distribution

of this sponge: (a) the minimum temperature at which reproduction can
occur; (b) the length of growing season which remains after the larvae have
settled, a factor which will determine the number of gemmules produced.
Halichondria bowerbanki, which has no winter-resistant gemmules, survives
the winter in inshore waters in abundance only during mild years. After severe
winters, recolonization on the inshore areas is largely dependent upon larvae
from colonies in deeper offshore waters.

Ill. SALINITY ‘TOLERANCE, COMPETITION FOR SUBSTRATE,
AND DISTRIBUTION AMONG CLIONIDS

INTRODUCTION

Although Long Island Sound opens into the Atlantic Ocean at both ends,
its western outlet is very narrow, preventing a rapid interchange of water between
the open ocean and the Sound, and the salinity is somewhat reduced. Galtsoff
and Loosanoff (1939) found that during August and September the bottom
salinity of the Sound varies gradually from 32 0/oo at its eastern end to 26 0/oo
near New York City. They reported that seasonal variations are small, the range
in any one locality seldom exceeding 1 to 2 o/oo during the year’s cycle. ‘The
difference in the salinity of water in the middle of the Sound from that of adja-
cent inshore localities is also slight, seldom more than 1 0/oo. According to a
recent survey made by Riley (1956), the annual range of salinities at stations in
the central part of the Sound is 4 to 5 o/oo.

The fauna of the greater part of Long Island Sound thus falls into the poly-
haline zone (16.5 o/oo to 30 0/oo) proposed in the scheme of classification of
brackish waters of Redeke (1922) as modified by Valikangas (1933). Remane
(1934) has doubted the validity of a polyhaline zone, pointing out that in waters
with salinities ranging down to 16.5 0/oo, there is simply a decrease in the num-
bers of marine species with few or no distinct brackish water species to be found.
Dahl (1948) has criticized Remane’s conclusions because of the latter’s use of
mean salinity values in a region (Kiel Bay) where great variations in salinity
occur. Dahl has given ample evidence of the existence of distinctive animal com-
munities living on algae in the polyhaline zone. Few, if any, of the species are
confined to this zone, but many have the center of their ranges here. He has
emphasized that in defining the lower limits of the marine zone, not only the
the mean value, but also the stability of the salinity factor is of importance. On
the basis of his studies of arthropods living on marine algae in the Skagerrak
and northern Cattegat, he was able to establish the lower limit of the marine
zone there at 27-28 o/oo, provided the variation coefficient is lower than 15. At
one station in the southern Cattegat, a mean salinity of 28 o/oo is combined with
a variation coefficient of 20, a range probably below that which a marine fauna
can endure. For a less stable area such as this, a value of 30-31 0/oo is a more
likely lower limit. Judging from the stable bottom salinity conditions found
by Galtsoff and Loosanoff and by Riley in Long Island Sound, this locality would
be an excellent one in which to study the polyhaline zone and its boundary with
the marine zone.

In the present study an attempt was made to determine the influence of
salinity tolerance on the distribution of five species of Long Island Sound sponges.
Work on three of these was soon abandoned because of difficulties in determining
death points. A satisfactory survival index was worked out for Cliona celata and
C. vastifica, however, and the results of salinity tolerance experiments on these
are reported, together with some observations on the distribution of these two
species and the possible importance of competition for substrate as a factor
in speciation.

106

SALINITY TOLERANCE 107

Previous experimental work on the salinity tolerances of sponges has been
limited to determining the effects of diluted sea water on reunition bodies. Galt-
soff (1925) found that dissociated cells of Microciona prolifera (Ellis and Solander)
failed to coalesce in hypotonic sea water with concentrations lower than 12.4 to
9.3 o/oo. He found that the cells are more sensitive to an increase in osmotic
pressure than to a decrease, coalescence taking place in hypertonic solutions up
to a value of 55.8 o/oo. De Laubenfels (1932a) found that dissociated cells of
Totrochota birotulata Higgin coalesced normally in solutions ranging from 80
per cent sea water (28 o/oo) to 110 per cent sea water (38.5 0/oo). Above and
below these values, the number of cell aggregates per unit area was lower.

SALINITY ‘TOLERANCES

METHops: Experiments to determine the salinity tolerances of Cliona celata
and Cliona vastifica were undertaken at the Biological Laboratory, U. S. Fish
and Wildlife Service, Milford, Connecticut, concurrently with similar experi-
ments on oysters being conducted by Dr. V. L. Loosanoff and his staff. Colonies
of the alpha-stage of Cliona celata and colonies of Cliona vastifica, both inhabit-
ing the shells of living Crassostrea virginica, were exposed for varying lengths
of time to salinities ranging from 27 o/oo to fresh water. The infected oysters
used were of approximately the same size and harbored colonies of sponges of
similar sizes. The oysters were collected on state leased oyster beds off Charles
Island, Milford, and in New Haven Harbor, at stations of essentially similar
depth and salinity characteristics.

The initial set of experiments was run by placing the oysters infected ‘with
sponge colonies in photographic trays supplied with running water of the fol-
lowing six concentrations: (1) 26-27 o/oo (Milford Harbor water, collected at
high tide), (2) 20 o/oo, (3) 15 0/oo, (4) 10 0/oo, (5) 5 o/oo, (6) fresh tap water.

The diluted concentrations were prepared in the following manner: A 5-liter
reservoir jar was kept filled with sea water up to the level of an overflow spout.
A similar jar was kept filled with fresh tap water. In this way a constant level
was maintained in each jar so that an equal pressure was applied at all times
to the outlet tubes leading from the bottom of each jar. To obtain a given diluted
concentration, an outlet tube from each reservoir jar was led into a beaker in
which the water had a chance to mix thoroughly before overflowing into the cul-
ture tray. A constant supply of a culture of “Chlorella” and miscellaneous flagel-
lates was introduced, dropwise, into the sea water reservoir.

The concentrations of water in the trays varied no more than 0.5 0/oo on
either side of the desired value. Hydrometric salinity determinations were made
twice daily to check this value.

After the initial experiments had demonstrated an approximate range for
survival, a set of less widely spaced concentrations was tried in the critical range,
mcluding the following two series: (1) 12 0/oo, 9 0/oo, 7.5 or 6 0/oo, 3 0/oo,
(2) 13 o/oo, 10 0/oo, 5 o/oo, 3 0/oo.

The experimental animals were exposed to the various concentrations of sea
water for periods of time ranging from one hour to one week and were in each
case transferred to sea water after exposure in order to note recovery. Observa-
tions were made during the course of and at the end of the period of exposure
to experimental conditions and also after a period of recovery in Milford Harbor
water (salinity, 26-27 0/oo).

Resutts: Under normal conditions Cliona celata bears ostia or incurrent pores

108 MARINE SPONGES

(from 5-10 in number) on the flattened summits of tubules which protrude from
holes in the surface of the calcareous substratum (PI. 1, fig. 2). Oscules are borne
individually on lower, wider tubules. Cliona vastifica, on the other hand, bears
both a single osculum and a group of ostia on every tubule.1 These structures
are illustrated by Volz (1939). The tubules in both species are capable of con-
traction into the inner recesses of the calcareous substratum so that the sponge
tissue is level with the surface of the substratum, and water currents through the
sponge cease. This fact provides the most readily observable criterion for cessa-
tion of activity in these organisms. Intermediate stages occur in which the tubules
are partially contracted and in which there is doubtless a reduced flow of water
through the animal. In some cases a few of the tubules were observed to be fully
extended with open oscules and pores, while others of the colony were in vari-
ous stages of contraction.

Upon prolonged exposure to lowered salinities, the tissue at the surface of the
sponge colonies turned brown; this state usually (but not invariably) preceded
failure of the colonies to resume normal activity within the period of observa-
tion. However, it is not certain whether such sponges were completely dead. In
some cases, breaking open the shell revealed living sponge tissue within, which
may have been capable of forming restitution bodies which could re-establish
the colony after the lapse of a sufficient amount of time in normal sea water.

Cliona colonies exposed to very low salinities (5 or 3 o/oo) and to fresh water
manifested a characteristic response. ‘The tubules contracted, in general, only
partially, and the tissue turned a pale, whitish color. This condition prob-
ably resulted from a rupture of the myocytes before complete contraction was
achieved.

The results of individual experiments are presented as histograms with the
aid of a survival index which summarizes the physiological condition of the col-
onies at the time of observation. Four categories of survival have been designated
as follows:

1. Colony inactive and seriously damaged. Tubules partly extended or com-
pletely contracted; oscules closed; color abnormal (brown or whitish).

2. Colony inactive. No flow of water through the sponge. Tubules partly
extended or completely contracted; oscules closed; color yellow.

3. Colony functioning, but at a reduced level. At least some of the tubules
extended or partly extended with some oscules and ostia open or partially open.

4. Colony functioning normally. Tubules extended; oscules and ostia open.

Considering first the experiments on Cliona celata, reference to the histograms
(figs. 34 through 39) which summarize the results reveal that this species is able
to function efficiently in waters with a salinity down to 20 o/oo. Exposure to
salinities as low as 15 0/oo causes some injury, but the sponge colonies are capa-
ble of carrying on at a reduced level of activity even after a week of exposure to
this lowered salt concentration (experiment 1D, fig. 36). In all cases complete
recovery followed a period of several days in sea water. In experiment IE, how-
ever, after a week of exposure to water of 15 o/oo, the colonies were inactive
and failed to recover after four days in sea water (fig. 36).

A salinity of 10 0/oo is near the critical one for Cliona celata. In experiment
1D, the colonies were still functioning at a reduced rate after 18 hours and 24
hours of exposure to 10 o/oo. These same colonies ceased functioning after
42 hours of exposure and were irreparably damaged after 72 hours of exposure to

1 Vosmaer (1933) describes separate incurrent and excurrent papillae.

SALINITY TOLERANCE 109

the same salinity. In all other cases in series 1, exposure to a salinity of 10 0/oo
(even for periods of two to six hours) led to cessation of activity in that environ-
ment, although recovery followed placement in sea water in the case of colonies
which had been kept in the lowered concentration for no longer than 24
hours.

In order to investigate tolerances in the critical range for survival, several
series of experiments were run using salinities ranging from 13 o0/oo to fresh
water. Series 2 consisted of four different exposures to salinities of 12, 9, 6, and
3 0/oo; series 3 involved exposures to concentrations of 12, 9, 7.5, and 3 0/oo;
in series 4, colonies were placed in salinities of 13, 10, 5, and 3 0/oo and in fresh
water. These experiments were carried out from September to December when
the water temperatures were decreasing steadily (fig. 41). The results, therefore,
are not strictly comparable to those of series 1 nor to one another.

In series 2, carried on in September when the water temperature was not ap-
preciably lower than it was during series 1, the sponge colonies were generally
inactive after one to 24 hours of exposure to the several lowered salinities (fig.
37). In a single instance partial functioning was recorded after 24 hours of ex-
posure to 9 o/oo. In general, complete or partial recovery followed transference
to sea water for several days, although sponges exposed to 3 o/oo for four hours
failed to resume a functional state after eight days in sea water, and colonies
which had been in concentrations of 6 and 3 o/oo for 24 hours also failed to
recover in sea water.

Series 3 was carried on in late October and early November, and results were
similar to those of the previous series. In no case was there evidence of water
currents flowing through the sponge colonies after periods of one to 96 hours of
exposure to salinities of 12, 9, 7.5, and 3 o/oo (figs. 38, 39). Complete recovery
followed removal to sea water after one, two, or four hours of exposure to the
lowered salinities and after 24 hours in salinities of 12 o/oo and 9 o/oo. In the
case of those colonies which were exposed to 7.5 and 3 o/oo for 24 hours, some
of the colonies recovered; others did not (three out of six in 7.5 0/00; two out
of six in 3 0/oo).

The lower temperatures under which the experiments in series 4 were carried
out produced a noticeable effect on survival. Experiments 4A and 4B (fig. 39)
are comparable to 3E in regard to length of exposure, and it is evident that in
the former instances some of the colonies remained active after being in the
lowered salinities for 42 and 48 hours. Similarly, recovery in sea water was greater
in series 4. The effect of lowered temperatures in increasing tolerances to lowered
salinities in Cliona celata is shown strikingly in figure 42 where mean survival
values after exposure and recovery in sea water are compared at 19° C. and
12.9°.G,

Tables 37 to 40 summarize the data of the salinity experiments on Cliona
celata and indicate along with figure 39 (series 4) the dependency of tolerance
to lowered salinities upon (1) length of exposure to the diluted concentrations
and (2) temperature.

Series 5 includes three experiments on the salinity tolerances of colonies of
Cliona vastifica (fig. 40). After 96 hours of exposure (5C) to salinities varying
from 27 o/oo to fresh water, complete recovery was noted after a week in sea
water for individuals exposed to salinities of 20 o/oo. Those which had been
immersed in water with a salinity value of 15 0/oo were not functional after a
week in sea water, but showed normal tissue within, so that eventual recovery

110 MARINE SPONGES

was probable. Those exposed to still lower salinities failed to show any signs of
recovery. In two sets of experiments (5A, 5B) carried on in the late fall, a sur-
prisingly high percentage of colonies survived exposure to even the lowest
salinities.

TABLE 37

SURVIVAL OF CLIONA CELATA AFTER VARYING PERIODS OF
EXPOSURE TO LOWERED SALINITIES—SrriEs 1

EXP. DURATION NO. OF CONCENTRATION OF SEA WATER IN PARTS PER MILLE
NO. OF (CLO) ODS RS
EXPOSURE 27 20 15 10 5 0
ee ee ee ei
Sa Oe ee nC ee
1D 18 hrs 3 4 4 3 3 1 1
1D 24 hrs 3 4 4 3 3 1 1
1D 42 hrs 2 4 4 3 2 2 1
1D 72 hrs 2 4 4 3 1 1 1
1E 72 hrs 3 4 4 cote 1 1 1
1D 7 days 1 4 4 3 1 1 1
1E 7 days 3 4 4 fas 1 1 | 1

* Numbers in these columns refer to survival index.
** Numbers in parentheses refer to number of colonies at each survival index level.

SALINITY TOLERANCE 111

TABLE 38

SURVIVAL OF CLIONA CELATA AFTER A 3-DAY PERIOD OF RECOVERY IN
SEA WATER FOLLOWING VARYING EXPOSURES TO LOWERED SALINITIES

SERIES 1
DURATION CONCENTRATIONS OF SEA WATER IN PARTS PER MILLE
EXP. OF NO. OF
NO. Tan UPTT nT |
EXPOSURE 27 20 15 10 5 0
3*(2)** | 3*(1)
1B 2 hrs 3 4 4 4 3 1 1
1C 6 hrs 3 4 4 4 3 1 1
1D 24 hrs. 1 4 4 4 1 1 1
1D UP Nive. 1 4 4 4 1 1 1
1D 7 days 1 4 4 4 1 1 1
1E 7 days 3 4 4 1 1 1 1

* Numbers in these columns refer to survival index.
** Numbers in parentheses refer to number of colonies at each survival index level.

MARINE SPONGES

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113

SALINITY TOLERANCE

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SURVIVAL
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>
i
2
3
=
WwW

20 %o

IS %o

10 %o

fresh
woter

AFTER
5 HRS.

RECOVERY RECOVERY
, 17 DAYS

3 3
(colonies) (cotonies)

EXPERIMENT IA
| hour exposure

RECOVERY
AFTER
5 DAYS
(3 colonies)

July 25-Aug. 11, 1947

EXPERIMENT 1B
2 hours exposure

SURVIVAL RECOVERY
AFTER AFTER
2 HRS. 3 DAYS
3 3
(colonies) (colonies)

RECOVERY
AFTER
3 HOURS
(3 colonies)

July 25-28, 1947

EXPERIMENT IC
6 hours exposure

SURVIVAL RECOVERY
AFTER AFTER
6 HRS 7 DAYS
3 3
(colonies) (cotonies)

RECOVERY
AFTER
3 DAYS
(3 colonies)

July 14-21, 1947

Figure 34. Summary of salinity experiments. Cliona celata, Series 1A, 1B, 1C.
Survival index explained in text, p. 108.

114

EXPERIMENT ID EXPERIMENT !0 EXPERIMENT ID
2; 18; 42 hours exposure 24 hours exposure 72 hours exposure

SURVIVAL 20 %o

INDEX
20%
15 %o
>
=
= Oe
=i
4
Ww
S%o
fresh
water
SURVIVAL SURVIVAL SURVIVAL RECOVERY SURVIVAL RECOVERY
AFTER AFTER AFTER AFTER AFTER AFTER
2 HOURS 42 HRS. 24 HRS. 6 DAYS 72 HRS. 4 DAYS
3 2 3 ! 2 '
(colonies) (cotonies) (cotonies) (colony) (cotonies) (colony)
RECOVERY RECOVERY
AFTER AFTER
RVIVAI RECOVERY
dlls ~ | HOUR 48 HRS. AFTER
18 HRS. i colony i colony 2 HRS.
3 colonies { colony
July 7-9, 1947 July 7-14, 1947 July 7-14, 1947

Ficure 35. Summary of salinity experiments. Cliona celata, Series 1D.
115

EXPERIMENT 1D
7 days exposure

SURVIVAL Qe
INDEX

20%o

IS %e

10 %o

SALINITY

G6G6@ OO

S%o

fresh
water

SURVIVAL RECOVERY

AFTER
7 DAYS

(colony)

OO

RECOVERY
AFTER
3 DAYS
(1 colony)

July 7-21, 1947

AFTER
7 DAYS

(cotony)

EXPERIMENT IE
72 hours; 7 days exposure

SURVIVAL RECOVERY
AFTER AFTER
72 HRS. 4 DAYS
3 3.
(colonies) (colonies)
SURVIVAL
AFTER
7 DAYS

(3 colonies)
July 14-25, 1947

FicurE 36. Summary of salinity experiments. Cliona celata, Series 1D, 1E.

SALINITY TOLERANCE 117

EXPERIMENT 2A EXPERIMENT 2B EXPERIMENT 2C EXPERIMENT 2D
| hour exposure 2 hours exposure 4 hours exposure 24 hours exposure

SALINITY

6 %o

3%o

SURVIVAL SURVIVAL SURVIVAL SURVIVAL
AFTER AFTER AFTER AFTER
| HOUR 2 HOURS 4 HOURS 24 HOURS
(2 colonies) (2 colonies) (2 colonies) (2 calonies)
RECOVERY RECOVERY RECOVERY RECOVERY
AFTER AFTER AFTER AFTER
8 DAYS 8 DAYS 8 DAYS 8 DAYS
(2 colonies) (2 colonies) (2 colonies) (2 colonies)
Sept. 10-18, 1947 Sept. 10-18, 1947 Sept.10-18, 1947 Sept. 4-10, (947

SURVIVAL
INDEX

Ficure 37. Summary of salinity experiments. Cliona celata, Series 2A through 2D.

118

EXPERIMENT 3A
t hour exposure

12%o

9%o
>
&
2
a)
<a
Ww
7.5 %o
3%o

VID

SURVIVAL
AFTER
' HOUR
(6 colonies)

D

RECOVERY
AFTER
5 DAYS
(6 colonies)

Oct. 3I-Nov. 5, 1947

MARINE SPONGES

EXPERIMENT 3B
2 hours exposure

a)
©

SURVIVAL SURVIVAL
AFTER AFTER
2 HOURS 4 HOURS
{6 colonies) (6 colonies)
RECOVERY RECOVERY
AFTER AFTER
6 DAYS 7 DAYS
(6 colonies) (6 colonies)

Oct. 30-Nov. 5, 1947 Oct.29-Nov. 5, 1947

SURVIVAL
INDEX

FicurE 38. Summary of salinity experiments. Cliona celata, Series 3A through 3D.

EXPERIMENT 3C
4 hours exposure

EXPERIMENT 3D
24 hours exposure

SURVIVAL
AFTER
24 HOURS
(6 colonies)

RECOVERY
AFTER
6 DAYS

(6 colonies)

Oct. 29-Nov. 5, 1947

SALINITY TOLERANCE 119

EXPERIMENT 3E EXPERIMENT 4A EXPERIMENT 4B
41,96 hours exposure 42 hours exposure 24 hours exposure

12 %o 13 %o

9%o 10 %o
>~
=
=
=I
ft
nn
>
Le
7.5 %o Se Oso
|
a
n
3 %o 3%o
fresh
SURVIVAL SURVIVAL
AFTER AFTER water
41 HRS. 96 HRS.
6 2
(colonies) (colonies)
RECOVERY RECOVERY
AFTER AFTER
5 DAYS 3 DAYS
(4 colonies) (2 colonies)
Oct. 27-Nov. 3,1947 SURVIVAL SURVIVAL
AFTER AFTER
42 HRS 48 HRS.
(6 colonies) (6 colonies)
RECOVERY RECOVERY
AFTER AFTER
5 DAYS 7 DAYS
SURVIVAL INDEX (6 colonies) (6 colonies)
‘ec. 8-15, 1947 Nov. 26-Dec. 5,1947

Ficure 39. Summary of salinity experiments. Cliona celata, Series 3E, 4A, 4B.

120 MARINE SPONGES

EXPERIMENT 5A EXPERIMENT 58 EXPERIMENT 5C
24 hours exposure 40 hours exposure 96 hours exposure

27 %o

20 %o

SALINITY

IS %o
10%
S%o
RECOVERY
AFTER
4 DAYS fresh
3 colonies water
Oct. 3I-Nov. 5, 1947
RECOVERY
AFTER
5 DAYS SURVIVAL RECOVERY
8 colonies AFTER AFTER
Nov.24-Dec. I, 1947 96 HRS. 7 DAYS
3 3
(colonies) colonies)
SURVIVAL
INDEX July (0-21, 1947

Ficure 40. Summary of salinity experiments. Cliona vastifica, Series 5A, 5B, 5C.

SALINITY TOLERANCE 121

DISCUSSION

OsMOREGULATION: An examination of the salinity determinations made twice
weekly during the year of 1947 at the Milford Biological Laboratory indicates
that variations in the salt concentrations of the bottom waters of Milford Harbor
were slight, ranging from 23.5 o/oo to 29 o/oo. The lowest records were made
following heavy rains, while the highest were recorded in late summer; the
mean is near 27 o/oo. Surface salinities show a much wider range of values,
varying from 5.2 0/oo after heavy rains to 29 o/oo in late summer. At an in-
termediate depth (three feet from the bottom in water six feet deep at mean low
water) the lowest salinity recorded over the course of a year was 12.2 o/oo. All
these records were made off the dock of the Milford Laboratory in water six feet
in depth at mean low water level and about 30 feet from shore line at the same
water level.

It is probable, therefore, that sponges inhabiting the bottom waters of Mil-
ford Harbor will seldom, if ever, come in contact with concentrations below 20
o/oo; so that the observations on salinity tolerances made in these experiments
suggest that the sponge colonies have wider tolerances than are likely to be met
with under natural conditions.

A capacity to adapt to wide changes in salinity may involve either (1) an active
regulation of osmotic pressure within the body which serves to insulate the tissues
from the salinity changes in the environment or (2) a wide tolerance to changes in
the osmotic pressure of the body fluids, with little regulative capacity. Within any
one phylum of invertebrates both conditions together with intermediates may
occur, so that a clear-cut differentiation into homoiosmotic and poikilosmotic
forms is impossible.

Since sponges have no body fluids, osmoregulation, if it exists, must be intra-
cellular; and a similarity to conditions in protozoans might be expected. Jepps
(1947) has shown that fresh-water sponges (genera Spongilla and Ephydatia) pos-
sess contractile vacuoles. Zeuthen (1939) found that the osmotic concentration
of the cells of Spongilla rises three- to fourfold at the time of gemmulation. Al-
though Jepps examined a variety of marine sponges, she was unable to find un-
equivocal evidence for the presence of contractile vacuoles in these forms. In
one case, however, Anchinoe (= Microciona) fictitia, clear fluid vacuoles were
observed in choanocytes. One such vacuole was seen to move forward during
the period of observation until it came to lie within the base of the collar, in the
position of the contractile vacuoles of the Spongillidae. Granule-containing
vacuoles were also observed moving in choanocytes, but in no case was either type
of vacuole scen contracting or discharging its contents. Although these vacuoles
may not have been rhythmically contractile, it is not unlikely that they do serve
to regulate water output. Kitching (1939) has pointed out that although marine
rhizopods lack contractile vacuoles, excess water introduced by food particles
is eliminated in food vacuoles. The vacuoles observed by Jepps in Anchinoe might
well function similarly.

True contractile vacuoles were reported in the choanocytes of Leucosolenia
by James-Clark (1868a, 1868b), but Jepps was unable to confirm these interesting
observations in her study of the same genus. Saville Kent (1880-82) figured con-
tractile vacuoles in Grantia compressa Bow., Leucosolenia coriacea Bow., and
Halisarca dujardini Johnston; however, he gave no statement in his text which
indicated that he had seen these structures functioning.

122 MARINE SPONGES

It may be suggested that sponges have been enabled to invade fresh waters
by virtue of the presence of contractile vacuoles which probably function as in
certain protozoans, as osmoregulatory mechanisms. The question of the possible
function of vacuoles in water relations in marine sponges which have succeeded
in entering brackish water remains to be further investigated. In marine sponges
without vacuoles it seems likely that extension of range in brackish waters depends
upon the tolerance of dilution of the internal milieu of the cellular elements, as
in the few coelenterates which have been studied. Concerning the coelenterates,
Bateman (1932) has critically reviewed the evidence for osmoregulation in the Scy-
phozoa and has presented work of his own showing that osmoregulation does
not exist in these forms, the concentration of salts in the tissues varying directly
with the external milieu. Palmhert (1933) studied the marine hydroid, Clava
multicornis (Forskal) and found no evidence for active osmoregulation. Exposure
to lowered salinities (down to 8-12 0/oo) resulted in an initial swelling of the
polyps followed by a slow return to normal size. Death followed 48 hours of
exposure to tap or pond water. Specimens placed in dilutions of sea water showed
a decreased oxygen consumption, another indication of an absence of osmoregula-
tory mechanisms. Miyawaki (1951) suggests that secreted mucus plays a role in
the resistance of actinians to low salinity. He found that Diadumene luciae (Ver-
rill) survived in sea water diluted to 7.5 per cent.

In multicellular animals with a body fluid, the environment of the tissue
cells can be regulated more readily. It has been shown by Wells and Leding-
ham (1940) that the rate of dilution is the determining factor in the sensitivity
of isolated polychaete muscle tissue to diluted sea water. Beadle (1937) demon-
strated that the tissues of intact polychaetes are protected from the effects of
sudden dilution of the sea water by a “damping” of the body fluid dilution curve
to the extent required for normal muscle functioning.

In a sense, a similar damping of the dilution curve of water in the canals of
a sponge could be brought about by the closure of the ostia and oscules by which
means a concentration of water higher than that in the environment would be
maintained in the channels of the sponge colony. Parker (1910), in his classical
experiments on the physiology of Stylotella (= Hymeniacidon) heliophila, found
that diluted sea water induced merely a partial and imperfect contraction of
ostia and oscules so that the internal cells were exposed to the lowered salinities
and activity ceased in dilutions of 50 per cent or lower after a period of 10-12
minutes. He also showed that upon exposure to fresh water, ostial and oscular
movements are paralyzed, and water currents cease immediately. Even after 24
minutes of such treatment, recovery occurs upon removal to sea water, the dam-
aged choanocytes being restored after several days.

In the present studies of Cliona a complete contraction of the oscules was
effected in the intermediate or critical salinities; that is, in salinities between
those in which some activity occurred (27-20 0/oo) and those in which immedi-
ate damage to the cells took place (5 o/oo—fresh water). This critical range of
salinities lies between 15 and 10 0/oo, and it is probable that a damping mecha-
nism analogous to that observed by Beadle may help sponges through limited
periods of exposure to these salinities through maintenance of a higher concen-
tration of internal water. Since most exposures of bottom-dwelling sponges to
fresh water under natural conditions (such as might occur after heavy storms
at inshore areas) would be preceded by a gradual lowering of the salinity, it is
probable that the sponge colonies would be completely contracted and thus pro-

SALINITY TOLERANCE 123

tected before the harmful concentrations are reached. Parker, however, found
that dilutions of sea water caused a relaxation of the ostia and oscules in
Hymeniacidon (Stylotella) if these structures were closed before placement in the
lower salinities; further investigations are needed to determine the extent to
which the oscules and ostia may function in protecting sponges from lower
salinities.

‘TEMPERATURE AND SALINITY TOLERANCE IN Gliona celata: In the investigations
of Cliona celata, experiments carried out under different temperature conditions
show distinctly different tolerances to diluted sea water. Figure 42 compares the
results of two comparable experiments. The mean survival and recovery values
for each concentration of diluted sea water are plotted for each temperature. In
experiment 3E, carried out in late October at temperatures averaging 19° C.,

TEMPERATURE — DEGREES CENTIGRADE

pss = NO On
TaSvra2CtmnnrsnGrvr =a2ZTa-CTHANPMAN
July Aug Sept Oct. Nov. Dec.

Figure 41. Water temperatures recorded during the salinity experiments. Each
point represents a weekly mean of the averages of fresh and sea water temperatures.

SALINITY Yoo (Series 3) SALINITY %o (Series 3)
12 9 75 3

x O—O Series 3-€ x

fo)
S Temp. 18.5 C.-19.5 ©. 2
=4 -=!
< @®--@ Series 4-A Ss
> >
« Temp. 12.3 C.—13.5 C. fea
>) 5
o 7)

13 10 DI: 3
SALINITY %o (Series 4) SALINITY %o (Series 4)

Ficure 42. Effect of temperature on tolerance to lowered salinities in Cliona celata.
Left, after 41 hours exposure to lowered salinities; right, after five-day recovery
period in sea water following the above exposures to lowered salinities.

124 MARINE SPONGES

the mean survival index after exposure varies from 1.5 to 1.0 and shows a similar
variation after “recovery.” In experiment 4A, carried out in late November when
temperatures averaged 12.9° C., the survival indices for recovery vary from 4.0
to 1.0, being from two to three times higher than in the previous experiment
except at a concentration of 3 o/oo, which was lethal in both cases.

These observations are in keeping with expectations since tolerance to re-
duced salinities involves physicochemical processes such as diffusion and _ per-
meability. Lucké and McCutcheon (1932), e.g., found a high effect of temperature
on the permeability of unfertilized sea urchin eggs, both in endosmosis and
exosmosis. They accounted for their observations in terms of changes in the vis-
cosity of water and in the permeability of the cell surface.

SALINITY TOLERANCE AS A FACTOR IN THE DISTRIBUTION AND SPECIATION IN THE
GENUs Cliona: When the experimentally determined tolerances of Cliona celata
and Cliona vastifica to reduced salinities are compared, the latter species is seen
to have a distinctly wider range of tolerance to this factor. Figure 43 shows the
percentage survival values of colonies of the two species after a 24 hour period of
exposure to a series of salinities varying from 12 0/oo to 3 o/oo, followed by a
recovery period of several days in sea water. The criteria for survival used were
(1) presence of open oscules with water currents issuing from them or (2) in the
event that all oscules were closed, survival was considered certain if the color
of the tissue was normal and if piles of shell chips could be detected at the base
of one or more of the oscules. Figure 43 also compares percentage survival values
of colonies of Cliona celata after recovery from 42 hours of exposure to salinities
varying from 13 0/oo to fresh water, with survival values of Cliona vastifica after
recovering from 40 hours of exposure to the same series of low salinities.

100 100

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S S
Bye z 75
= =)
w w
w Ww
q 50 © 50
e bE
a a
rs) rs)
oes 25
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3 6 2 l2 @—® C celata

SALINITY-%o O--O © vastifica SALINITY—%o

Ficure 43. Comparative tolerance to lowered salinities of Cliona celata and C.
vastifica. Left, after 24 hours of exposure to lowered salinities; right, after 40 hours
of exposure to lowered salinities.

In the case of both sets of data (24 hours and 40 hours of exposure respectively)
straight lines have been fitted to the points by the method of least squares. The
regression coefficients for the two species in each case differ significantly at a level
lower than one per cent.

This experimental evidence suggests that salinity tolerance may play a role
in determining the distribution of these two species and that salinity may have
operated as an isolating factor in speciation.

A consideration of the world-wide occurrence of the two species of Cliona
concerned reveals that their distributions overlap in large areas, and that the

SALINITY TOLERANCE 125

relative abundance of the two species varies considerably. Cliona vastifica occu-
pies a greater variety of habitats, ranging from low water neap tides to depths
of 600 meters. It is also found consistently in waters of reduced salinity where
Cliona celata is excluded. The latter species ranges from mean low water to
depths of 200 meters (rarely). Further consideration of the general distribution
of these two species with a discussion of competition between them will be found
in a later section. For the present, the occurrence of these species in brackish
waters will be treated specifically.

The limitations placed upon the distribution of C. celata and C. vastifica by
reduced salinity are parallel over a large part of their range: both occur in the
mouth of the Chesapeake Bay, in Beaufort Harbor, in Long Island Sound, in the
Belt Seas of Denmark, in the Etang de Thau. In Louisiana the data of Hopkins
(1956a) indicate that C. celata is found farther up estuaries than is vastifica; how-
ever, only three colonies of the latter species (2 per cent of all clionids identified)
were found during his studies. None were recorded from waters of higher salinity
(his zone 5). In South Carolina, too, Hopkins (1956b) found that vastifica makes
up a relatively small proportion of the total clionid population (9.5 per cent of all
boring sponges identified, as compared to celata which comprises 74 per cent of the
total). C. celata, alone, was found at station 4 (South Santee River) where the
lowest salinity (16.6 0/oo) of his survey was recorded. A salinity of 26 o0/oo was
the lowest observed at stations where C. vastifica was found. The salinity records
of Hopkins’ South Carolina survey have a limited significance, however, since
they represent single determinations in most instances.

One striking fact stands out in reviewing the distribution of Cliona vastifica.
Populations which seem to bear a close relationship to vastifica in morphological
traits have penetrated into brackish (mesohaline) waters in four regions of the
world: the Atlantic and Gulf coasts of North America, the Oued Melah, the
Black Sea, and Chilka Lake. The first form, which is sufficiently differentiated
from C. vastifica to merit specific rank, has been named C. truitti? by Old (1941).
The seaward limit of its range in Chesapeake Bay occurs in a region where the
surface isohalines show an annual range of 15 to 17 o/oo and the annual range
of salinities at 20 meters is 14 to 20 0/oo (Cowles, 1931). The extension of its
range into the headwaters of the bay includes waters with a surface salinity
down to 6 o/oo and an annual range at 20 meters of 10 to 16 o/oo. It occupies
a region approximately equivalent to Valikangas’ pleiomesohaline zone. The
boundary between the area inhabited by C. truztts and that occupied by C. vas-
tifica and C. celata is very sharp according to Old’s data (fig. 44).

Hopkins (1956a, 1956b) has studied the distribution of Cliona truitti: on the
coasts of Louisiana and South Carolina and has presented excellent salinity data
for the waters where collections were made in the former locality. Hopkins has
used the clionids as “salinity indicators” and has attempted to classify the estua-
rine waters of Louisiana into six “Cliona zones.” His observations show that
the range of C. truitt? in Louisiana overlaps those of celata, vastifica, and lobata
(see Table 41), unlike the distribution patterns given by Old for Chesapeake Bay.
Only in Hopkins’ zone 1, with annual salinity range and mean values of 1-11-25
o/oo, and with salinities below 5 0/oo recorded on 15 per cent of the days of the
year, does Cliona truitti occur to the exclusion of all other clionids. Cliona celata
is found along with truitti in zones 2 through 5.

2 Old also reports C. truitti from Long Island Sound but gives no exact locality. It
would be of interest to know whether it is restricted to brackish water bays there.

39° 30

Baltimore

DELAWARE
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397

38° 30°

Sinnepuxent |/ A

Chincoteague
Bay

38°

A
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CHESAPEAKE

37° 30°

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® = Cliona vastifica
28%. ‘A Cliona celata

37°
James R.

Norfolk Cape

¢ Henr
C2 Wa y

76° 30° 76° 75° 30° 75°
Ficure 44. Distribution of clionids in the Chesapeake Bay region in relation to
salinity.
126

127

SALINITY TOLERANCE

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128 MARINE SPONGES

Hopkins’ South Carolina studies (1956b) are of great interest. An accumulated
rainfall deficiency of about 47 inches in the last seven years combined to a lesser
extent with a recent rise in sea level in that area have led to an increased salinity
in the estuaries. “Sea water had replaced brackish water of low salinity in most of
the estuaries; water of almost oceanic salinity was found even in small creeks far
inland.” Where possible, salinity data recorded by Lunz during a survey of coastal
waters of South Carolina in 1935 (reported by Hopkins, 1956b) are compared
with the 1956 salinity records of Hopkins in ‘Table 42. Although based in all cases

TABLE 42
SALINITY RECORDS IN SOUTH CAROLINA ESTUARIES IN 1935 AND 1956

LOCALITY SALINITY IN 1935 (LUNZ)| SALINITY IN 1956 (HOPKINS)
Rantowles Creek 9.1 0/oo 31 0/oo
Five Fathom Creek (mouth) 14.0, 17.8, 19.7 0/oo 30.6 0/00
Wadmalaw River 1971; 20:0, 21'.5'6/c0 32 0/oo
Fish Creek (mouth at St. 32.6 0/00 32 0/oo
Helena Sd.) (middle of St. Helena Sd.)
Ashepoo River 25.6, 27.0, 29.6 0/oo 29 o/oo

(1-4 mi. above 1935 record)

Mackay and Skull Creek 26.9, 28.7, 29.3 0/oo 26.2-31.2 0/oo (1955)

on single salinity determinations these data give some idea of the magnitude of
the changes in salinity which have occurred at some stations.

In regard to the distribution of clionids in South Carolina, Hopkins found
that C. truztit is relatively much less abundant there than in Louisiana. In South
Carolina truitti comprised 10 per cent of the 1527 boring sponges identified by
Hopkins, whereas 73 per cent of the 148 specimens identified in Louisiana were
truitti. These facts suggest that the invasion of South Carolina estuaries with
water of higher salinity is resulting in a replacement of Cliona truitti by the
other three species of Cliona found there. This conclusion is borne out especially
well by the data from Rantowles Creek, in which 1950 and 1956 collections are
compared (Table 43). Salinity records for Rantowles Creek in 1950 are not given,
but available data indicate a sharp rise in salinity since 1935: 9.1 o/oo, 1935;
17.6 o/oo, 1955; 31.0 0/oo, 1956.

No experimental data are available as yet in regard to the tolerance of
C. truitti to higher salinities. ‘Thus it is difficult to say at present whether this
species is dying back in South Carolina because it is unable to tolerate more
saline waters which are moving up the estuaries or because of competition for
substratum or food with the other clionids which are now able to live farther up
the rivers.

SALINITY TOLERANCE 129

TABLE 43

RELATIVE ABUNDANCE OF CLIONIDS AT RANTOWLES CREEK IN 1950
AND 1956

NUMBER OF EACH SPECIES IDENTIFIED AND PER CENT OF TOTAL

YEAR ——

Cliona celata Cliona truattt Cliona vastifica Cliona lobata
1950 17p@z) 8 (89%) 0 0
1956 49 (38%) 39 (30%) 4 (3%) 38 (29%)

Old’s collections of Cliona robusta and C. spirilla® were made at Piver’s Is-
land, Beaufort Harbor, N. C., where salinity variations have been studied by
Gutsell (1931). The ranges and means of the monthly maximum and minimum
salinities as observed by Gutsell at Piver’s Island over a four year period are
38-33-24 o/oo and 35-25-6 o/oo, respectively. The low reading of 6 0/oo was
made after a heavy rain in September; minimum salinities of 14 to 20 o/oo are
not infrequent in the late fall and winter months. De Laubenfels (1947) recorded
salinities as low as 15 o/oo in July, 1946, during an exceptionally rainy summer.
No information is available about the occurrence of clionids in Newport River
where the salinity values are lower.

In the Black Sea the degree of difference of the local form (described as
C. stationis) from typical specimens of C. vastifica as it occurs in the Mediter-
ranean is not as great. The surface salinity of the Black Sea is 18 0/oo, increasing
somewhat with depth (Nikitin, 1931), and it may be that the clionid population
is not completely isolated from that of the Mediterranean proper. In Chilka Lake
the local form is barely distinguishable from the typical open sea variety.

In all these cases the oxeas and spirasters are similarly modified in waters of
lowered salinities, although the extent of modification varies greatly. The acan-
thoxeas tend to be longer and to possess central swellings, this characteristic being
most pronounced in the case of C. truitti. This centrotylote condition is merely
an intensification of a tendency present in Cliona vastifica in general. Many of
the acanthoxeas in specimens of this species from Long Island Sound observed
by the present writer have a trace of centrotylosis. The spirasters tend to be
shorter in the brackish water forms and are, in general, less angulated than in
typical specimens of Cliona vastifica. In addition to spicule differences, there
is a noticeable enlargement of the sizes of oscular and ostial tubules in the case
of C. truitti as compared with C. vastifica. Information in this regard is lacking
for the other brackish water types.

Topsent (1932) has reported an abnormal variety of Cliona vastifica collected
by Seurat in brackish water in the Oued Melah at Nador on the coast of ‘Tunis.
Specimens of the same species of sponge from a station in the open sea at nearby
Skira show a typical structure. The tylostyles of the brackish water form are
little modified. The diactinal megascleres are smooth oxeas, contrary to any other
brackish water populations in the world; but these oxeas do tend to have a

3 Hopkins (1956b) has found specimens in South Carolina intermediate between Cliona
spirilla and vastifica and suggests that they may be conspecific.

130 MARINE SPONGES

central swelling, which seems to be a consistent development in all areas of
greatly reduced salinity. The microscleres are noteworthy for their sparse spina-
tion, unusual thickness, and a tendency to be straight rather than spiral. A few
of the microscleres are longer than the rest and tend to be pointed at one end
as though they were intermediate between spirasters and oxeas. Czerniavsky
(1880) described a species, Cliona pontica (from the Black Sea coast of the Cau-
casus at Sukhumi in water 1.5 to 15 meters deep) with smooth oxeas, many of
which are figured as being centrotylote in form. No spirasters were found in
these specimens. Nasonov (1925) was unable to find specimens fitting the descrip-
tion of C. pontica among Czerniavsky’s collections from Sukhumi in the Zoologi-
cal Museum of the Academy of Sciences of the U.S.S.R. He did find Cliona lobata
in the shells of these collections, however. Nikitin (1934) recorded only Cliona
stationis during the course of his survey of the Gudaut oyster bank near Sukhumi.

It is of interest to note in passing that microscleres seem to be more sensitive
than megascleres to changes in the chemical conditions of the environment. This is
obvious from the present information in regard to the influence of lowered sa-
linity, and it was found to be equally true by Jewell (1935) and J¢rgensen (1944)
who raised fresh water sponges in water of reduced silicon content.

It would be instructive to acclimatize colonies of Cliona vastifica to waters of
lowered salinities in order to determine whether modifications of the spicules
such as are outlined above are environmentally induced. Similarly, C. truztti
should be raised in waters of higher salinity to find out whether the modifications
observed are genetically stabilized. Spicule sizes of the forms considered above
are summarized in Table 44.

The experimentally demonstrated salinity tolerances of C. celata and C. vas-
tifica reveal that both species are capable of functioning in environments with
salinities lower than those ordinarily met with in their range of occurrence.
Both are capable of withstanding very low salinities for short periods, a protec-
tive capacity necessary for any organism which penetrates inlets with variable
salt concentrations. C. vastifica has been shown to possess the capacity of func-
tioning in lower salinities than is true of C. celata. This, as will be apparent
from the more detailed analysis of its geographical distribution, is merely an-
other instance of its generally higher tolerance of environmental extremes. It
is not surprising therefore, that C. vastifica should have an initial advantage
under conditions of lowered salinity such as have led to the formation of Cliona
truittt in Chesapeake Bay and of the brackish water populations of C. vastifica
elsewhere in the world. It is strange that C. vastifica has not penetrated more
deeply into the Baltic Sea; perhaps this fact is correlated with the restricted east-
ward range of Ostrea edulis although certainly other molluscs are available for
colonization there. The fact that the Baltic has been connected with the North
Sea for a relatively short time may explain the absence of a distinct population
of C. vastifica there.

The gradation in differentiation of the brackish water populations of Cliona
vastifica throughout the world is a matter of great interest. There is a range of
variation from full species formation and probable reproductive isolation in
Chesapeake Bay to subspecies formation in the Black Sea and incipient varia-
tion in Chilka Lake, and finally, to the complete absence of a brackish water
population in the Baltic Sea. If adequate studies of the microfossils in the sedi-
ments of all these regions were available to provide information on the history
of the salinity changes of these bodies of water, some concept of the time taken

SALINITY TOLERANCE 131

TABLE 44
SPICULE SIZES OF CLIONA VASTIFICA AND RELATED FORMS

SPECIES LOCALITY, TYLOSTYLES | ACANTHOXEAS SPIRASTERS
AUTHOR Lx WwW Lx W Lx W
Cliona vastifica Atlantic Coast 160-1504 50-110yu 6-23
Old, 1941 x 3-5yu x 2.5-4u x 1-3.5y
Cliona truttts Chesapeake Bay | 190-225 110-130u 7-12u
Old, 1941 5-3. 50 x 2.5-3.5u x 0.5-2u
Cliona vastifica Rovigno 280-306 95-132u 11-16y
Volz, 1939 x 3.5-5.5u x 3-4 x 1.5-3u
Cliona vastifica Sevastopol 168-340 120-140 8-12u x 2y
Swartschewsky, | x 3-9u x 2-3u
1905
Cliona stationis Sevastopol 245-313 yu 75-175p. 9-13u
Nasonoy, 1925 x 5-7 x 2.5-10yu
(at center)
Cliona vastifica Oued Melah, 260-290 (smooth oxeas)} 9—20yu, usually
Tunis x 4u 55-110u 12-15y, x 2.5-4p
Topsent, 1932 (7.54 at base) | x 3-5u (some 30—40u x 4p)
Cliona robusta Beaufort Harbor | 160-200u (spined or 8-16 x 1-2u
Old, 1941 x 2-4 smooth oxeas)
40-130u
x 4-12u
Chiona spirilla Beaufort Harbor | 160—230u 80-170u 9-20u x 1-2y
Old, 1941 x 2-4u x 3-9u

for speciation under the differing conditions of temperature and availability
of substratum could be obtained. More extensive biometric studies of the sponge
populations in each case are also needed.

THE ZOOGEOGRAPHY OF CLIONA CELATA, CLIONA VASTIFICA,
AND OTHER CLIONIDS

INTRODUCTION: Animals boring into calcareous matter, which is limited in
amount in any environment, might be expected to be in strong competition
for substratum. The substratum factor represents an area of overlap in the defi-
nition of the niches of all such animals in any one region. Differences in the
ecology of such animals can be sought in feeding mechanisms, intrinsic growth
rate, and breeding periods. ‘The summary of the distributions of Cliona celata
and Cliona vastifica given here, demonstrates that these species coexist over a large
part of their range, but that one or the other of them is invariably more abun-

132 MARINE SPONGES

dant. All of the substratum available is probably never occupied; on Long Island
Sound oyster beds, for instance, about 80 per cent of the oysters are infected with
boring sponge colonies. Hopkins (1956b) found that 72 per cent of the 2116
shells which he examined in South Carolina were infected with clionids. Simi-
larly it is doubtful if every available bit of substratum of the limestone cliffs
of the Adriatic Sea is occupied. Nonetheless, it is apparent that a competitive rela-
tionship exists from the frequent occurrence of colonies of two species in the
same oyster shell in the former locality and by the restricted and specialized
substratum requirements exihibited by some of the species found in the latter
region.

The number of species of clionids which coexist in any locality varies con-
siderably as a function of the amount of substratum available, a fact which is
apparent from Table 45. Warm limestone areas seem to be most favorable for

TABLE 45
OCCURRENCES OF CLIONIDAE ON DIFFERENT SUBSTRATA

LOCALITY NUMBER OF SPECIES DOMINANT AUTHOR
SUBSTRATUM

Rovigno d’Istria 9(3)* Limestone Volz, 1939
Beaufort, N. C. 5(2) Oyster shells Old, 1941
Chesapeake Bay 4(1) Oyster shells Old, 1941

Long Island Sound 4 Oyster shells Old, 1941
West-central Pacific 5(1) Dead coral de Laubenfels, 1954
Bermuda 2(1) Corals de Laubenfels, 1950
Gulf of Maine | 1 Scallop shells Hartman

(Deep water)

* Endemic species in parentheses.

the growth and speciation of clionids; brackish water oyster beds provide an
abundance of substratum and favorable conditions for speciation. Coral reefs
do not present as wide a variety of clionids as might be expected. De Laubenfels
(1950, 1954) reports two species from Bermuda and five from the west-central
Pacific. The latter species show little overlap in distribution, however. Cliona
lobata Hancock and C. vastifica Hancock occupy distinct atolls in the Marshall
Islands; C. lobata was also recorded from Truk and C. euryphylla Topsent from
Ponapé in the eastern Carolines; C. schmidti (Ridley) was found in the Palaus;
Aka trachys de Laubenfels is described by its author as possibly the only clionid
in the Marianas. Burton (1934) reports no clionids from the Great Barrier Reef
although he believes that two species of Spirastrella there may pass through a
boring stage in early life.

DISTRIBUTION IN TEMPERATE REGIONS: A detailed tabulation of the occurrences

SALINITY TOLERANCE 133

of Cliona celata and Cliona vastifica is unnecessary in the present work since
both Vosmaer (1933) and Volz (1939) have recently compiled such records. In-
stead, attention will be focused on several well studied areas in an attempt to
ascertain the limiting factors in the distribution of these two species.

Along the Atlantic Coast of North America, the distribution of these species
has been studied by Lambe (1896), George and Wilson (1919), Procter (1933),
Old (1941), de Laubenfels (1947, 1949), Hopkins (1956a, 1956b), and the present
writer. Cliona celata occurs in inshore waters from Prince Edward Island to the
coast of South Carolina, reaching its greatest abundance in waters of somewhat
lowered salinity (30 to 25 0/oo) such as occur in coastal bays and sounds where
oysters are cultivated extensively. This species is especially common in Malpeaque
Bay (Prince Edward Island), Buzzard’s Bay, Narragansett Bay, Long Island Sound,
Delaware Bay, Chesapeake Bay (southern part), Beaufort Harbor (N. C.), and
in the estuaries of the South Carolina coast. In the Gulf of Mexico it has been
reported from St. George’s Sound, Florida (Menzel, 1956), from oyster beds along
the coast of Louisiana (as Cliona sulphurea by Moore, 1899, and Cary, 1906a, b;
1907a, b; also by Hopkins, 1956a), and has been identified by the present writer
in oyster shells from Aransas Bay, Texas. Hopkins (1956a) also reports celata from
Texas bays.

Cliona vastifica also ranges widely along the American Atlantic Coast. It has
been collected by the present author in Pecten magellanicus shells in the Bay
of Fundy (off Grand Manan Island, 30 meters), in a fragment of an unidentified
mollusc in the Gulf of Maine (Fippennies Ledge, 80 meters), and in the shells
of Pecten magellanicus in Massachusetts Bay (Stellwagen Banks, 40 meters). It is
noticeably absent from inshore waters in northern areas. Procter (1933) failed
to find C. vastifica in his survey of the Mt. Desert Island fauna, although C. celata
is common there; de Laubenfels does not record it from Woods Hole where
C. celata is again the common boring sponge. In Long Island Sound Cliona vas-
tifica occurs along with C. celata on oyster beds. In the experience of the present
writer, C. celata is about twice as common as C. vastifica on oysters near New
Haven, Connecticut. The percentage of infection of Crassostrea virginica shells
by boring sponges in a sample collected on an oyster bed off Woodmont, Con-
necticut, is given in Tables 46 and 47.

Old (1941) has recorded C. vastifica from the southern shores of Long Island
Sound but has given no indication of its abundance relative to C. celata. C. vas-
tifica was also found by Old in York River, Chesapeake Bay, and in Beaufort
Harbor, N. C. Lunz (1935) and Hopkins (1956b) recorded it on oyster beds along
the South Carolina coast where it made up 9.5 per cent of all clionids examined
by Hopkins. Careful search would probably reveal its presence in Delaware Bay,
Narragansett Bay, and Buzzard’s Bay. Pearse and Wharton (1938) listed it from
Apalachicola Bay, Florida; Hopkins (1956a) recorded it from Louisiana where it
comprised 2 per cent of clionids identified.

Thus Cliona celata is the dominant species in inshore waters along our coast.
It is apparently capable of occupying a larger proportion of the calcareous sub-
stratum necessary for its early life than the other species (chiefly C. vastifica, C.
truittt, and C. lobata) living sympatrically with it. On offshore scallop grounds,
however, Cliona vastifica is the exclusive or at least the dominant boring sponge.
Its incidence of occurrence on living scallop shells is much lower than the total
infection of oyster shells by boring sponges. Between 75 and 80 per cent of all
oysters in the New Haven region of Long Island Sound contain boring sponges;

134 MARINE SPONGES
TABLE 46

NUMBER OF LIVING OYSTERS INFECTED BY BORING SPONGES IN
LONG ISLAND SOUND

SPECIES OF SPONGE INFECTING OYSTERS NUMBER OF OYSTERS PERCENTAGE OF

INFECTED TOTAL

Cliona celata 81 40.5 \ ane
Cliona vastifica 46 23.0
Cliona celata and Cliona vastifica 14 7.0
Cliona lobata 7 355
Cliona celata (Dead colonies) 10 5.0
Sponge colonies absent 42 21.0
TOTAL 200 100.0

TABLE 47

NUMBER OF SHELLS OF DEAD OYSTERS INFECTED BY BORING SPONGES
IN LONG ISLAND SOUND

SPECIES OF SPONGE INFECTING SHELLS NUMBER OF SHELLS INFECTED

Living Cliona celata colonies 14
Dead C. celata colonies* 19
Total number of C. celata colonies 33
Living C. vastifica colonies 5)
Dead C. vastifica colonies* 16
Total number of C. vastifica colonies 21
Shells with both species, C. celata and C. vastifica 7
Cliona lobata 4
Shells lacking sponge colonies 35
TOTAL 100

* The clionid galleries in some of these shells may have been excavated by sponges while the
oysters were still alive and before the shells were thrown back in the water as cultch.

SALINITY TOLERANCE 135

whereas the incidence of infection of scallops by clionids in Massachusetts Bay
and the Gulf of Maine is of the order of 25 per cent. The higher rate of infection
of oysters is doubtless partly explicable in terms of the tendency of these molluscs
to clump on old shells. In this way a single sponge colony can spread asexually
to six or seven oysters.

It is probable that the faster growth rate of C. celata, which is manifest in
its eventual overgrowth of the shells it infects to form a free- living colony, €x-
plains the greater abundance of this form over all other species in certain regions.
Cliona vastifica grows more slowly; seldom, if ever, forms free-living colonies;
and seems to be incapable of fully exploiting the ecological niche which is avail-
able to it on the scallop beds. The absence of C. celata from the Massachusetts
Bay scallop beds may result from an incapacity to produce larvae in the cooler off-
shore waters with a consequent dependence upon inshore stocks to colonize
deeper areas. Since oyster beds north of Cape Cod are largely depleted, the boring
sponge cannot attain the abundance which it does south of the Cape, and the like-
lihood of its establishment in offshore waters from inshore-produced larvae is
slight. Its absence from the northern part of the Gulf of Maine and from the Bay
of Fundy is consistent with its range in inshore waters in the North Atlantic.
Its northernmost recorded occurrence in North America is in Malpeaque Bay,
Prince Edward Isl.; it is absent from Greenland (Bréndsted, 1914, 1916, 1933a,
1933b). In the eastern North Atlantic it occurs at the Shetland Isls. (Bowerbank,
1882) and in Stavanger Bay, Norway (Burton, 1930a); it is absent from northern
Norway (Burton, 1930a), the Faroes (Br¢ndsted, 1932), and the White Sea (Mere}-
kowsky, 1879; Swartschewsky, 1906). This distribution suggests that the tempera-
ture for larval production is the limiting factor in its northern distribution since
it is perennial in growth along the Atlantic Coast of the United States where
winter temperatures equal those of Arctic regions. Vertically, its distribution
extends down to 200 meters and, to the present writer’s knowledge, its occurrence
in deep waters is restricted to tropical areas except for one record at 130 meters
off the west coast of France where the mean annual temperature is about 15° C.

The occurrence of Cliona celata and Cliona vastifica in the North and Baltic
Seas is shown in figure 45. Cliona celata is widely distributed on the oyster beds
of Holland, the East and North Frisian Islands, Denmark, and Schleswig-Hol-
stein (Arndt, 1935, 1943). Levinsen (1893) records C. celata in the Skagerrak and
Cattegat at depths of 18-32 meters. So far as present records indicate, Cliona
vastifica is largely absent from this extensive region. One specimen has been
reported from the deepest part of the Kieler Bucht (Arndt, 1933), and there are
several records of its occurrence on the Lophohelia banks off the Gullmars and
Vaderé Fjords and in the Koster area, all in southwestern Sweden (Fristedt, 1885;
Alander, 1942). Arndt (1943) records it from @resund and from the North Sea
north of the 60 meter line. The Swedish records are from depths of 40 meters or
over. (Such depths occur very close to shore in this region contrary to the case
along the Danish coast which slopes gradually.) Cliona celata is reported by Fri-
stedt and Alander as being common in the shallow coastal waters of Sweden.
Alander has suggested that the restriction of C. vastifica to deep waters in the
Swedish area may be a result of its incapacity to withstand the low salinities of
the surface waters; however, the salinities in this region vary from 30 to 20 o/oo
(Valikangas, 1933) and it is unlikely that such would prove lethal to this species.
Bottom salinities in the Kieler Bucht vary from 20 to 15 o/oo (Schulz, 1932).

Nasonov (1925) has reported Cliona vastifica from the Barents Sea along the

136 MARINE SPONGES

B Be es :
Stavanger
Boyf-

NORWAY

‘\
NORTH SEA
\

-
-——

aoam

NORTH
FRISIAN
IS.

~=
-_—
OP ee eas aaa

~ HELGOLAND oh
.* EAST FRISIAN IS.

/ Ache 7 mh

if weeseer
WEST FRISIAN Is.
o ID = se i

Surface salinities

Feb. eeceeees Aug: — —

6°

BALTIC SEA

GERMANY

fi Cliona vastifica
A Cliona celata

12°

Ficure 45. Distribution of clionids in the North Sea area in relation to salinity.

northern shore of the Kola Peninsula. It was found at a depth of 140 meters in
the Kola Gulf near the eastern shore of Sedlovaty Island and in shallow water in

the Bay of Oura of the Motovka Gulf.

DISTRIBUTION IN THE MEDITERRANEAN ReEcION: The Adriatic Sea is especially
well known as a result of the work of Lendenfeld (1898) and Volz (1939). The
latter author in particular has made very careful studies of the ecological relation-
ships of the nine species of the family Clionidae in the region of Rovigno d’Istria.
His observations are summarized in figure 46; see also Hartman (1957).

SALINITY TOLERANCE 137

Volz reports the greatest abundance of clionids in the littoral terrace extend-
ing from low water springs to a depth of 25 meters. Here seven species coexist
with Cliona celata and C. vastifica, which are abundant only in shallow water and
again in deep water. Throughout most of the terrace these two species tend to
occur chiefly in mollusc shells and assume an insignificant role compared to
C. viridis and C. vermifera and other dwellers in limestone.

In the deeper waters below the limestone terrace is an area of sand and shells
which supports three species of boring sponges; Cliona vastifica, Cliona viridis,
and Cliona celata, in order of abundance. C. vastifica occurs almost exclusively
in mollusc shells, especially those of mussels. C. viridis, on the other hand, occurs
mainly in association with coralline algae. Cliona celata is a less common inhabi-
tant of both substrata.

In addition, Cliona vastifica lives in the narrow zone between low tide neaps
and low tide springs, and is the only species to inhabit the limestone cliff at this
level. Thus, in summary, it may be said that C. vastifica is the most widespread

| HIGH TIDE| NEAPS
pe (et LOW TIDE] NEAPS

CLIONA ALBICANS
CLIONA SCHMIDT!

CLIONA VERMIFERA
CLIOTHOSA HANCOCKI
THOOSA MOLLIS
CLIONA ROVIGNENSIS

CLIONA VASTIFICA
CLIONA VIRIDIS

CLIONA CELATA
SOVees” AVEO

suajaw SZ Of

suajow Se—Sl
SNOZ TW3HS OSNTIOW GNV GNVS

VERY COMMON

COMMON

MODERATELY COMMON

UNCOMMON

UE &

Ficure 46. Vertical distribution of clionids at Rovigno. Data after Volz.

138 MARINE SPONGES

species in the Rovigno district, occurring from low water neaps down to the deep
offshore waters. In the two extreme environments it is the dominant species; in
the intermediate littoral terrace it is pushed into an insignificant place by com-
petition with the fast-growing C. viridis and C. vermifera. C. celata is a minor
species in this part of the Adriatic, although it ranges throughout the littoral
terrace and into the sand and shell zone, and also is able to survive in tidal pools
above mean low water when it is thrown up into such locations by the action
of the surf. It does not occur on the oyster beds in the canal of Leme where the
salinity is reduced, although C. vastifica, C. viridis, and C. vermifera are found
there in limited numbers.

Several other Mediterranean records of C. celata and C. vastifica are of in-
terest in the present connection. Seurat (1934) noted that C. vastifica alone occurs
in the zone between low water neaps and low water springs in the Gulf of Gabés,
thus extending the observations of Volz. Both species occur in the Etang de Thau,
a brackish water inlet on the south coast of France (Topsent, 1925b).

The most frequently recorded boring sponge in the Black Sea was described
by Nasonov (1883) as a new species, Cliona stationis. It has been reported from
the Bay of Sevastopol (Nasonov, 1883, 1924; Swartschewsky, 1905; Kudelin, 1910),
from the Bay of Odessa (Kudelin, 1910), and from Sukhumi (Nikitin, 1934). Ac
cording to Nasonov’s figures (1883) the spicules are highly variable in form.
Many of the tylostyles are characterized by irregular knobs and swellings along
their lengths, and the oxeas, which are usually spined but may be smooth, show
a tendency toward centrotylosis. The spirasters are rodlike to contorted in shape,
and some have central swellings. Swartschewsky (1905) figured quite normal tylo-
styles and acanthoxeas, the latter without central swellings, and has synonymized
C. stationis with C. vastifica. Kudelin (1910) also reports this sponge as C. vastifica.
Nikitin (1934) retains the name C. stationis. It is apparent that the vastifica-like
clionids of the Black Sea are highly variable in spiculation and are probably
best regarded as a subspecies, Cliona vastifica stationis (Nasonov). Czerniavsky’s
(1880) species, C. pontica, is of uncertain status, as mentioned earlier, but it is
possible that it represents an extreme variant of vastifica.

FURTHER DISTRIBUTION: Annandale (1915a) states that Cliona vastifica is the
commonest species in the littoral zone of the east coast of India. It often makes
its way into brackish waters in this region, occurring in Chilka Lake at Orissa
and the Ganjam district (in shells of Ostrea and Purpura), in the Adyar River
at Madras, and in the Ennur Backwater (in Ostrea shells). It is common in the
Persian Gulf in shells of Avicula and Margaritifera; it occurs in Placuna shells
from Palk Straits (514 fathoms) and in the shells of Oliva and Malleus from the
Andaman Islands. Cliona celata is not nearly as common in Indian waters, only
three records being given by Annandale. These are from a station 28 fathoms in
depth off the coast of Burma (in calcareous algae); from shallow water at Madras
(in Pyrula); and from 614 fathoms in the Gulf of Manaar (also in Pyrula).

In Chilka Lake, Cliona vastifica is very abundant on the oyster beds of the
outer channel (Annandale, 1915b). The salinity of this part of the lake is about
as high as that of the open sea (ca. 35 o/oo) during eight months of the year, but
in December flood waters reduce the salinity to values between 17.4 and 5.3 o/oo.
A condition of almost fresh water is maintained for the next three months. Liv-
ing specimens of Cliona vastifica have been taken from Chilka Lake when the
salinities were 8.9 and 14.1 o/oo. The change back to normal sea water often

SALINITY TOLERANCE 139

occurs in the course of a single day. Although C. vastifica is most abundant in the
Ostrea shells of the outer channel, it also occurs in the main area of the lake
where the salinity is always reduced; in this area it inhabits the shells of Purpura
(Thats) carinifera.

SUMMARY OF THE ZOOGEOGRAPHY OF THE CLIONIDAE: Let us examine the zoogeo-
graphical data in an attempt to determine some of the factors which have led to
speciation in the family Clionidae. In the first place, the Mediterranean data tend
to confirm the generally held idea that the center of speciation of this family of
sponges is in the warm waters where calcareous matter abounds. Here in shallow
waters nine species of clionids coexist. Coral reefs have been considered as the
most probable sites of origin of boring sponges, and in this regard it might be
mentioned that the spirastrellids which grow in cavities in corals are suggestive
of early stages in the development of the excavating habit. Both Vosmaer (1911)
and Burton (1934) have noted species of the genus Spirastrella which are restricted
in occurrence to cavities in corals. It is uncertain whether or not the sponge colony
has helped in forming the cavities; certainly they are not nearly as complex as the
galleries actively excavated by clionids. But the obligatory relationship of some
spirastrellids with this type of habitat suggests a stage in the development of the
boring process. Morphologically, Spzrastrella is closely allied to Cliona; indeed,
the most diagnostic character separating them is the boring habit of the latter.

Extension of range into temperate regions has depended upon an abundance
of mollusc shells such as occur in coastal regions where Ostrea abounds or in
offshore areas where Pecten lives.

If the discussion is restricted now to Cliona celata and Cliona vastifica, it is
found that the overall ranges are practically coextensive, but that local occur-
rences are variable in the well-studied areas which have been cited as examples.
C. celata is the dominant form on the oyster beds of the American Atlantic Coast
and in the North Sea and Belt seas of Western Europe. In the first locality Cliona
vastifica occurs sympatrically with C. celata at least as far north as Long Island
Sound, but is only about half as abundant as the first-named species. On western
European oyster beds (at least those in the North Sea) Cliona celata is the only
species found. On both sides of the Atlantic in temperate regions, the range of
C. vastifica extends into deeper offshore waters: the scallop areas of Massachusetts
Bay and the Gulf of Maine; the Lophohelia banks off southwestern Sweden; the
southern part of the Barents Sea; and the deepest parts of the Kieler Bucht. In
these areas it is without competition from other clionids for substratum. Where
oysters are especially common both species can find sufficient substratum to co-
exist; under such conditions C. celata predominates in temperate waters. Where
oysters are less common (as in the inshore waters of the Gulf of Maine) present
evidence indicates that Cliona celata is the sole inhabitant of these shells, and
C. vastifica has exploited offshore areas.

In the warmer regions of the world, Cliona vastifica is more common than
C. celata. In the Adriatic, e.g., C. vastifica occurs from the low water neap tide
level down to the deep sand and shell areas. It is the sole clionid in the restricted
area between low water neaps and low water springs; but it fares only moderately
well in the multispecific competition for substratum which occurs in the littoral
terrace. Here two other species (Cliona viridis and Cliona vermifera) are dominant
and six other species occur in the same area. At least two of these occupy strictly
delimited niches. Cliona vastifica tends to prefer mollusc shells although it is also
found in the limestone cliff; Cliona celata is common in the littoral terrace in

140 MARINE SPONGES

shells and in limestone. Finally in the deeper areas, C. vastifica assumes the domi-
nant role again, inhabiting the mollusc shells which are abundant there. C. viridis
is less common and occurs chiefly in coralline algae; C. celata is of uncommon
occurrence in both shells and algae.

Along the coasts of India, Cliona vastifica is again the abundant form with
C. celata of less common occurrence.

Cliona vastifica is, in the final analysis, a more adaptive and plastic species,
being capable of inhabiting a wider variety of habitats in regard to depth, reduced
salinity, and exposure to air than is true of C. celata. A factor of importance in the
wider tolerances of C. vastifica is doubtless the capacity to form gemmules in which
state it can live through periods of stress. Cliona celata is better adapted to ex-
ploit such habitats as the oyster beds of temperate regions by virtue of its faster
growth rate, larger size, and probable concomitant reproductive advantages. How-
ever, along with its specializations which enable it to exploit a particular habitat,
is a restricted adaptability to environmental extremes. Thus, it does not occur on
the scallop beds, possibly because it cannot reproduce at the low temperatures
present at such depths.

In the warm waters of the Adriatic the dominating position of C. celata dis-
appears before that of two other species which seem better adapted to existence in
limestone. Still stranger is the fact that in Indian waters C. vastifica is dominant
over C. celata on oyster beds. These facts suggest that C. vastifica is the older
species and that C. celata arose in temperate regions where it is well adapted to
exploit the beds of gregarious molluscs occurring there. It has since spread into
warmer seas, but always occupies a minor role in the competition for substratum
in these localities. Even on the Indian oyster beds it does not assume the dominant
position which it occupies under similar circumstances in temperate waters. Young
specimens of C. celata (in Europe at least) often possess oxeas and spirasters, sug-
gesting an origin from a species like C. vastifica with a full complement of spicules.
The fact that young specimens of C. celata with microscleres have seldom been
recorded from American waters suggests that this species developed in the Old
World and subsequently spread to the American continent.

A NOTE ON THE CONTROL OF BORING SPONGES

Since the clionids infect such a high percentage of the oysters in Long Island
Sound and other inlets along the North American Coast, and since their excava-
tions make the shells of oysters weak so as to hinder processing in canning fac-
tories, it is of considerable interest to oystermen in these areas to develop control
measures. Topsent (1900) suggested that immersion for a short time in fresh water
would suffice to kill the boring sponges in oysters. De Laubenfels (1947) dipped
an infected oyster “briefly” in fresh water and noted that the boring sponge colony
failed to recover in sea water. The present experiments indicate that at tempera-
tures near 23° C. a period of exposure to fresh water between one and two hours
would be necessary to kill the sponge colonies. Even then it is quite probable that
restitution bodies would arise from the cells which were hidden in the recesses of
the excavations and eventually regeneration would ensue. In experiment 1A of the
present investigations it was found that although recovery from one hour of ex-
posure to fresh water was slow, after a period of 17 days in sea water, the sponges
were fully functional once again. Under the conditions of oyster farming prac-
ticed in America, exposure of the animals to fresh water for several hours is
hardly feasible. If a substance toxic to the sponge colonies were found it is con-

SALINITY TOLERANCE 141

ceivable that it could be effectively applied in solution to piles of oysters on the
decks of boats.

Another approach to the problem concerns the type of cultch used. Normally
dead oyster and clam shells are planted out on the beds to serve as substrata for
the attachment of spat. This cultch is planted out in late spring in time for the
most intensive setting period of oysters in July. The setting period of boring
sponges is in August and September so that this calcareous cultch is subject to
infection the same year in which it is planted out. Since a single shell often sup-
ports six to eight young oysters, it is obvious that during the following year of
growth the sponges can infect all of these animals by asexual spreading. It is likely
that the high incidence of infection among oysters as compared to scallops is a
result of this very fact. Hence it is suggested here that a non-calcareous cultch
such as broken bricks or tiles would decrease the incidence of clionid infection by
isolating more of the oysters from their neighbors and thus preventing asexual
spreading of the colonies. Perhaps coating the cultch shells with black varnish
would protect them from infection by boring sponges; this method was suggested
by Orton (1937) as a means of darkening shells to attract more spat.

Additional information is needed concerning the intensity of settling of clionid
larvae. The work reported here from Milford Harbor indicated a rather low rate
of infection by this means; however, conditions may be quite different on the
oyster beds at New Haven Harbor where clionids reach a peak of abundance.

SUMMARY

1. Both Cliona celata and C. vastifica are capable of functioning normally in
salinities lower (20 0/oo) than they are likely to meet in offshore waters in Long
Island Sound.

2. Both species can recover to normal activity after brief exposures to salinities
down to 10 0/oo (or lower).

3. Tolerance of clionids to low salinities is inversely related to temperature.

4. Cliona vastifica is more tolerant of exposure to low salinities than C. celata.

5. Populations of Cliona vastifica have invaded brackish water in a number of
regions of the world and have undergone parallel morphological changes in each
area, although the extent of differentiation varies. Three populations along the
American Atlantic Coast apparently merit specific rank.

6. The zoogeography of the family Clionidae is discussed. It is found that
Cliona celata is the dominant species on oyster beds in temperate regions; Cliona
vastifica dominates on Indian oyster beds. In each case the other species assumes a
secondary position in the same area and in some cases occurs as the dominant
form in adjacent offshore areas.

7. Cliona celata probably originated in the Old World in temperate regions
and has since spread to tropical areas where it has difficulty in invading niches
already filled. The generally increased size and higher growth rate of Cliona celata
together with higher reproductive capacity have favored its establishment in in-
shore temperate regions, but in warmer areas other unknown factors operate in
keeping it in a secondary position.

8. It is suggested that the use of a non-calcareous cultch for collecting oyster
spat would reduce the incidence of infection of oysters by boring sponges.

REFERENCES CITED

ALANDER, HARALD

1942. Sponges from the Swedish west coast
and adjacent waters. Goteborg, Henrik
Struves, 95p., 16 pl.

ALLEE, W. C.

1923. Studies in marine ecology: I. The dis-
tribution of common littoral inverte-
brates of the Woods Hole region. Biol.
Bull. Woods Hole, vol. 44, 167-191, 1 pl.

ANNANDALE, NELSON

1915a. Indian boring sponges of the family
Clionidae. Rec. Indian Mus., vol. 11 (1),
124 ee ie ae

1915b. Fauna of the Chilka Lake: Sponges.
Mem. Indian Mus., vol. 5, 23-54, 11 fig.,
Pl. 3-5.

ARNDT, WALTHER

1933. Ein fiir die Kieler Bucht und die Ostsee
neuer Bohrschwamm. Schr. naturw. Ver.
Schl.-Holst., vol. 20, 54-55.

1935. Porifera. Tierwelt N.-u. Ostsee, Teil

IIa, Lief. 27, 140p., 239 fig.

Eine neuere Ausbeute von Meeressch-

wammen der West- und Siidkiiste Portu-

gals. Mem. Mus. zool. Univ. Coimbra,

(1) no. 116, 75p., 2 fig.

1943. Die tiergeographische Gliederung der
Schwammfauna der Nord- und Ostsee.
Arch. Naturgesch., (2) vol. 12, 303-349.

1941.

ARNESEN, EMILY

1917. Brutknospenbildung bei Polymastia
mammillaris (O. F. Miiller) Bow. K.
norske vidensk. Selsk. Skr., 1917 (1), 24
p., 1 fig., 6 pl.

BATEMAN, J. B.

1932. The osmotic properties of medusae. J.
exp. Biol., vol. 9: 124-127.

BATTARRA, J. A.

1773. Rerum naturalium historia nempe quad-
rupem insectorum piscium variorum-
que marinorum corporum _fossilium
plantarum exoticarum ac_ praesertim
testaceorum existentium in Museo Kir-
cheriano .. . Edited by A. P. Philippo
Bonannio. Rome, Aere Venantii Mo-
naldini Bibliopolae.

BAUHINO, JOHANNE AND J. H. CHERLERO

1651. Historia plantarum universalis nova et
absolutissima: cum consensu et dissensu

circa eas. Ebroduni, vol. 3, 866p.

BEADLE, L. C.

1937. Adaptation to changes of salinity in the
polychaetes. I. Control of body volume
and of body fluid concentration in
Nereis diversicolor. J. exp. Biol., vol. 14,
56-70.

BERGMANN, WERNER

1949. Comparative biochemical studies on the
lipids of marine invertebrates, with spe-
cial reference to the sterols. J. Mar. Res.,
vol. 8 (2), 137-176, 2 fig.

Brwper, G. P.

1928. Some sponges of the south-west coast.
Proc. Sthwest. Nat. Un. 1927, 12-20,
2 fig.

BiGELow, H. B.

1914. Explorations in the Gulf of Maine, July
and August, 1912, by the U. S. Fisheries
Schooner Grampus. Oceanography and
notes on the plankton. Bull. Mus. comp.
Zool. Harv. vol. 58 (2), 29-147, 38 fig.,
9 pl.
Physical oceanography of the Gulf of
Maine. Bull. U. S. Bur. Fish., 1924, vol.
40 (2), 511-1027, 207 fig.
1933. Studies of the waters on the continental
shelf, Cape Cod to Chesapeake Bay. I.
The cycle of temperature. Pap. phys.
Oceanogr., vol. 2 (4), 1-135, 66 fig.

1928.

BoWERBANK, J. S.

1859. On the anatomy and physiology of the
Spongiadae. Part I. On the spicula. Phil.
Trans. roy. Soc. London, vol. 148, 279-
332, Pl 23-20:

In: MacAndrew, Robert, “List of the
British marine invertebrate fauna.” Rep.
Brit. Ass. Adv. Sci., vol. 30, 217-236.
On the anatomy and physiology of the
Spongiadae. Part III. On the generic
characters, the specific characters, and
on the method of examination. Phil.
Trans. roy. Soc. London, 1862, vol. 152,
1087-1135, Pl. 72-74.

A monograph of the British Spongiadae.
Ray Soc. Publ., London, vol. 1, 290p.,
37 pl.

A monograph of the British Spongiadae.
Ray Soc. Publ., London, vol. 2, 388p.
A monograph of the British Spongiadae.
Ray Soc. Publ., London, vol. 3, 367p.,
O2 ple

A monograph of the British Spongiadae.
Edited by A. M. Norman. Ray Soc.
Publ., London, vol. 4, 250p., 17 pl.

1861.

1863.

1864.

1866.

1874.

1882.

REFERENCES CITED

BR@NDSTED, H. V.

1914. Conspectus Faunae Groenlandicae. Po-
rifera. Medd. Grgnland, vol. 23, 459-544.

1916. Report on the Porifera collected by
the Danmark Expedition at North-East
Greenland. Danmark-Eksped. til Grgnl.
Nordgstkyst, 1906-1908, vol. 3 (17), 475-
483, Pl. 25-26.

1932. Marine Spongia, in: Zoology of the Fa-
Foes, vol. 3, 54p., Il fig:

1933a. The Godthaab Expedition, 1928. Porif-
era. Medd. Grgnland, vol. 79 (5), 25p.,
8 fig.

1933b. The Scoresby Sound Committee’s Sec-
ond East Greenland Expedition in 1932
to King Christian IX’s Land. Porifera.
Medd. Grgnland, vol. 104 (12), 10p.

BurTON, MAURICE

1926. Observations on some British species of
sponges belonging to the genus Reniera.
Ann. Mag. nat. Hist., (9) vol. 17, 415—
aoe

1928. A comparative study of the character-
istics of shallow-water and deep-sea
sponges, with notes on their external
form and reproduction. J. Quekett micr.
Cl., vol. 16, 49-70, 7 fig., Pl. 3.

1930a. Norwegian sponges from the Norman
Collection. Proc. zool. Soc. Lond., 1930,
487-546, 8 fig., Pl. 1-2.

1930b. Additions to the sponge fauna at Ply-
mouth. J. mar. biol. Ass. U.K., (2) vol.
16, 489-507, 9 fig.

1932. Report on a collection of sponges made

in South Saghalin by Mr. ‘Tomoe Urita.

Sci. Rep. Tohoku Imp. Univ., (4) Biol.

vol. 7 (2), 195-206, 6 fig., Pl. 7-8.

Sponges. Sci. Rep. Gr. Barrier Reef

Exped. 1928-1929, vol. 4 (14), 513-621,

33 fig., 2 pl.

Some sponges from the Okhotsk Sea and

the Sea of Japan. Explor. Mers russes,

vol. 22, 61-81, 6 fig.

Notes on sponges from South Africa,

with descriptions of new species. Ann.

Mag. nat. Hist., (10) vol. 17, 141-147,

4 fig.

Notes on the ecology of sponges. Brit.

Sci. News, vol. 2 (15), 83-85, 4 fig.

Suberites domuncula (Olivi): its syn-

onymy, distribution, and ecology. Bull.

Brit. Mus. (nat. Hist.) Zool., vol. 1 (12),

353-378.

CarTER, H., J.

1880. In: D’Urban, W. S. M., “The Zoology
of Barents Sea.” Ann. Mag. nat. Hist.
(5)) vol. 6G? 253-277.

1882. Some sponges from the West Indies and
Acapulco in the Liverpool Free Mu-
seum described, with general and classi-

1934.

1935.

1936.

1949.

1953.

143

ficatory remarks. Ann. Mag. nat. Hist.,
(5) vol. 9, 266-301, Pl. 11-12.
Catalogue of marine sponges collected by
Mr. Jos. Willcox on the West Coast of
Florida. Proc. Acad. nat. Sci. Philad.,
1884, 202-209.

Gary, E.R:

1906a. The conditions for oyster culture in the
waters of the parishes of Vermilion and
Iberia, Louisiana. Bull. Gulf biol. Sta.,
vol. 4, 1-27, 1 chart.

1906b. A contribution to the fauna of the
coast of Louisiana. Bull. Gulf biol. Sta.,
vol. 6, 50-59.

1907a. The cultivation of oysters in Louisiana.
Bull. Gulf biol. Sta., vol. 8, 1-56, 6 pl.

1907b. A preliminary study of the conditions
for oyster culture in the waters of Terre-
bonne Parish, Louisiana. Bull. Gulf biol.
Sta., vol. 9, 1-62.

CELESIA, PAOLO

1893. Della “Suberites domuncula” e della
sua simbiosi coi Paguri. Boll. Mus. Zool.
Anat. comp. Genova, No. 14, 63p., Pl.
5-8.

Cog, W. R.

1932:

1885.

Season of attachment and rate of growth
of sedentary marine organisms at the
pier of the Scripps Institution of Ocean-
ography, La Jolla, Calif. Bull. Scripps
Instn Oceanogr. tech., vol. 3 (3), 37-86,
3 fig., Pl. 1-6.

Core, W. R. AND W. E. ALLEN

1937. Growth of sedentary marine organisms
on experimental blocks and plates for
nine successive years at the pier of the
Scripps Institution of Oceanography.
Bull. Scripps Instn Oceanogr. tech., vol.
4 (4), 101-136, 6 fig., Pl. 1-2.

Corte, JULES

1914. L’association de Cliona viridis (Schmidt)
[Spong.] et de Lithophyllum expansum
(Philippi) [Algues]. C. R. Soc. Biol.,
Paris, vol. 76, 739-740.

Coues, ELLIOTT, AND H. C, YARROW

1879. Notes on the natural history of Fort
Macon, N. C., and vicinity. (No. 5) Proc.
Acad. nat. Sci. Philad., 1878, 297-315.

Cow es, R. P.

1931. A biological study of the offshore waters
of Chesapeake Bay. Bull. U. S. Bur.
Fish., 1930, vol. 46, 277-381, 16 fig.

CZERNIAVSKY, VOLDEMARO

1880. Spongiae littorales Pontis Euxini et
Maris Caspii. Bull. Soc. Nat. Moscou,

1879, vol. 54 (2), 88-128, 288-320, PI.
5-8.

144

DAHL, ERIK

1948. On the smaller Arthropoda of marine
algae, especially in the polyhaline wa-
ters off the Swedish west coast. Under-
sdkningar Over Oresund, vol. 35, 193p.,
42 fig.

DEEVEY, E. S., Jr.

1947. Life tables for natural populations of
animals. Quart. Rev. Biol., vol. 22 (4),
283-314, 9 fig.

De LAUBENFELS, M. W.

1932a. Physiology and morphology of Porifera
exemplified by Jotrochota birotulata
Higgin. Publ. Carneg. Instn, no. 435,
37-66, 6 fig., 2 pl. (Pap. Tortugas Lab.
vol. 28).

1932b. The marine and fresh-water sponges of
California. Proc. U. S. nat. Mus., vol.
81 (4), 1-140, 79 fig.

1936. A discussion of the sponge fauna of the
Dry Tortugas in particular and the
West Indies in general, with material
for a revision of the families and orders
of the Porifera. Publ. Carneg. Instn,
no. 467, 225p., 22 pl., (Pap. Tortugas
Lab. vol. 30).

1947. Ecology of the sponges of a brackish
water environment at Beaufort, N. C.
Ecol. Monogr., vol. 17, 31-46, 2 fig.

1949. The sponges of Woods Hole and ad-

jacent waters. Bull. Mus. comp. Zool.
Harv., vol. 103 (1), 1-55, Pl. 1-3.

The Porifera of the Bermuda Archi-
pelago. Trans. zool. Soc. Lond., vol. 27
(i) d= 154, 65 fig 2 pie

The sponges of the west-central Pacific.
Ore. St. Monogr. Zool., No. 7, 306p.,
200 fig., 12 pl.

Sponges of Onotoa. Pacif. Sci., vol. 9
@);, 137-143; 2 fig:

DENDY, ARTHUR

1950.

1954.

1955.

1914. Observations on the gametogenesis of
Grantia compressa. Quart. J. micr. Sci.,
(2) vol. 60, 313-376, Pl. 23-26.

1924. Porifera. Part I. Non-Antarctic Sponges.
Nat. Hist. Rep. Terra Nova Exped.
1910. Zool., vol. 6 (3), 269-392, 15 pl.

DENDY, ARTHUR AND L. M. FREDERICK

1924. Porifera. Rep. Canad. arct. Exped. 1913-
1918. vol. 8 (J), 8p.

DeEsor, EDOUARD

1851. On two new species of sponges. Proc.

Boston Soc, nat. Hist., 1848, vol. 3,

67-68.

Dusoscg, O. AND O. TUZET

1937. L’ovogénése, la fécondation, et les pre-
miers stades du développement des

MARINE SPONGES

éponges calcaires. Arch Zool. exp. gén.,
vol. 79, 157-316, 42 fig., Pl. 6-19.

EHLERS, E.

1870. Die Esper’schen Spongien in der zoo-
logischen Sammlung der K. Universitat
Erlangen. Erlangen. 36 p.

ELLIs, JOHN

1755. An essay towards a natural history of
the corallines and other marine prod-
ucts of the like kind commonly found
on the coasts of Great Britain and Ire-
land. London, 103p., 39 pl.

1786. The natural history of many curious
and uncommon zoophytes, collected
from various parts of the globe. (Edited
by Daniel Solander) London, Benj.
White and Son, 206p., 63 pl.

Esper; E. J. G.

1794. Die Pflanzenthiere in Abbildungen nach
der Natur mit Farben erleuchtet nebst

Beschreibungen. Zweyter Theil. Niirn-
berg. 303 p., 49 pl.

FLEMING, JOHN

1828. History of British animals. Edinburgh,
Bell and Bradfute, 565p.

FRISTEDT, KONRAD

1885. Bidrag til kinnedomen om de vid Sve-
riges vestra Kust lefvande Spongiae. K.
svenska VetenskAkad. Handl. vol. 21
(6), 56p., 4 pl.

Sponges from the Atlantic and Arctic
Oceans and the Behring Sea. Vega-
Exped. Vetensk. Iakttag., vol. 4, 401-

471; Ble 22-31.

1887.

FUGLISTER, F. C.

1947. Average monthly sea surface tempera-
tures of the western North Atlantic
Ocean. Pap. phys. Oceangr., vol. 10 (2),
25p., 2 fie.) 16"pl:

GALTsoFF, P. S.
1925. Regeneration after dissociation (an ex-
perimental study on sponges). 1. Behav-
ior of dissociated cells of Microciona
prolifera under normal and altered con-
ditions. J. exp. Zool., vol. 42, 183-221,
10 fig., 1 pl.

GaLtsorr, P. S. AND LOOSANOFF, V. L.

1939. Natural history and method of con-
trolling the starfish (Asterias forbesi,
Desor). Bull. U. S. Bur. Fish., vol. 49
(31); 75-132; 32 he:

GEORGE, W. C. AND H. V. WILSON

1919. Sponges of Beaufort (N. C.) Harbor and
vicinity. Bull, U. S. Bur. Fish., 1917-
1918, vol. 36, 129-179, Pl. 56-66.

REFERENCES CITED

GINNANI, G.

1757. Opera postume. Vol. 1. Venice. (Not
seen by the writer.)

GRANT, R. E.

1825. Observations and experiments on the
structure and functions of the sponge.
Edinb. phil. J., vol. 13, 94-107, 333-346.

1826a. Observations and experiments on the
structure and functions of the sponge.
Edinb. phil. J., vol. 14, 113-124.

1826b. Notice of a new zoophyte (Cliona celata
Gr.) from the Frith of Forth. Edinb.
new phil. J., Apr.-Oct. 1826, 78-81.

1826c. Observations on the structure of some
silicious sponges. Edinb. new phil. J.,
Apr.-Oct. 1826, 341-351.

1827. Observations on the structure and func-
tions of the sponge. Edinb. new phil. J.,
Oct. 1826-Apr. 1827, 121-141, Pl. 2.

1841. Outlines of comparative anatomy. Lon-
don, Hippolyte Bailliere, 656p., 147 fig.

Gray, J. E.

1867. Notes on the arrangement of sponges,

with the description of some new gen-
era. Proc. zool. Soc. London, 1867, 492-
558, Pl. 27, 28.

GRENTZENBERG, MAX

1891. Die Spongienfauna der Ostsee. Inaugu-
ral-Dissert. Univ. Kiel., 54p., 38 fig.

GUTSELL, J. S.

1931. Natural history of the bay scallop. Bull.
U. S. Bur, Fish., 1930, vol. 46, 569-632,
32 fig.

Hancock, ALBANY

1849. On the excavating powers of certain
sponges belonging to the genus Cliona;
with descriptions of several new species,
and an allied generic form. Ann. Mag.
nat... Hust; |(2)¢vol<3, 321-348, PI 12-15:
Note on the excavating sponges; with
descriptions of four new species. Ann.
Mag. nat. Hist. (3) vol. 19, 229-242, PI.
1-8.

1867.

HANITSCH, RICHARD
1890. Third report on the Porifera of the

L.M.B.C, District. Proc, Lpool biol. Soc.
vol. 4, 192-233, PL. 10-15,

1891. Notes on some sponges collected by Pro-
fessor Herdman off the west coast of Ire-
land from the ‘“‘Argo.” Proc. Lpool biol.
Soc: vol. 5, 203-2227) 2 i

1894. Revision of the generic nomenclature

and classification in Bowerbank’s “Brit-
ish Spongiadae.” Proc. Lpool biol. Soc.,
vol. 8, 173-206.

145

Hartman, W. D.

1957. Ecological niche differentiation in the
boring sponges (Clionidae). Evolution,
vol. 11 (3), 294-297, 1 fig.

HARTMEYER, ROBERT

1914. On Alcyonium pulmonis instar lobatum
Ellis. J. Mar. biol. Ass. U. K., (2) vol.
10, 262-282, 1 fig.

HENTSCHEL, ERNST

1929. Die Kiesel- und Hornschwimme des
Nordlichen Eismeers. Fauna arct., Jena,
vol. 5 (4), 857-1042, Pl. 12-14.

HERLANT-MEEWIS, HENRIETTE

1949. Contribution a l'étude histologique des
Spongiaires. Ann. Soc. zool. Belg., vol.
79; 9-90,,7 Be:

Hopkins, S. H.

1956a. Notes on the boring sponges in Gulf
Coast estuaries and their relation to
salinity. Bull. mar. Sci. Gulf Carib.
vol. 6, 44-58.

1956b. The boring sponges which attack South
Carolina oysters, with notes on some
associated organisms. Contrib. Bears
Bluff Labs., no. 23, 30p. 1 map.

Hutcuins, L. W. AND MARGARET SCHARFF

1947, Maximum and minimum monthly mean
sea surface temperatures charted from
the “World Atlas of Sea Surface Tem-
peratures.” J. Mar. Res., vol. 6 (3), 264-
268, fig. 69, Pl. 1, 2.

IMPERATO, FERRANTE

1599. Dell’ historia naturale di Ferrante Im-
perato Napolitano. Naples, Constantino
Vitale. 791p.

JAMEs-CLARK, H.

1868a. An investigation of the nature of
sponges. Proc. Boston Soc. nat. Hist., vol.
11, 16-17.

1868b. On the Spongiae ciliatae as Infusoria
flagellata; or observations on the struc-
ture, animality, and relationship of
Leucosolenia botryoides Bowerbank.
Ann. Mag. nat. Hist., (4) vol. 1, 133-
ba S82 bee 250-264, ble oa

Jepes, M. W.

1947. Contribution to the study of the
sponges. Proc. roy. Soc. London, vol.
B 134, 408-417, 5 fig.

JEWELL, M. E.

1935. An ecological study of the fresh-water
sponges of northern Wisconsin. Ecol.
Monogr., vol. 5, 461-504, 26 figs.

146

JORGENSEN, C. B.

1944. On the spicule-formation of Spongilla
lacustris (L.) I. The dependence of the
spicule-formation on the content of dis-
solved and solid silicic acid of the mi-
lieu. K. danske vidensk. Selsk. Biol.
Medd., vol. 19 (7), 45p., 16 figs.

JOHNSTON, GEORGE

1842. A history of British sponges and litho-
phytes. Edinburgh, W. H. Lizars. 264p.,
25) pl:

JORGENSEN, O. M.

1917. Reproduction in Grantia compressa.
Rep. Dove Mar. Lab., (2) vol. 6, 26-32,
1 pl.

1918. Note on the larvae of Grantia com-
pressa. Rep. Dove Mar. Lab., (2) vol. 7,
60-61, 1 pl.

KENT, W. S.

1880-82. A manual of the Infusoria. 3 vols.,
London, D. Bogue, 913p., 51 pl.

KINGSLEY, J. S.

1901. Preliminary catalogue of the marine In-
vertebrata of Casco Bay, Me. Proc. Port-

land Soc. nat. Hist., vol. 2, 159-183.

KITCHING, J. A.

1939. The physiology of contractile vacuoles.
IV. A note on the sources of the water
evacuated, and on the function of con-
tractile vacuoles in marine Protozoa.
J. exp. Biol., vol. 16, 34-37.

KUuDELIN, N.

1910. Zur Frage der Spongien des Schwarzen
Meeres. Mém. Soc. Nat. nouv.- russ. Est.,
vol. 35, 1-40, Pl. 1-2.

LAMARCK, J. B. P.

1815. Suite des polypiers empatés. Mém. Mus.
Hist. nat., Paris. vol. 1, 162-168.

LAMBE, L. M.

1892. On some sponges from the Pacific Coast
of Canada and Behring Sea. Trans. roy.
Sac. Ganz, vol: 10 (4); 67-78; Pl. 3-6:
Sponges from the Pacific Coast of Can-
ada. Trans. roy. Soc. Can., vol. 11 (4),
Patras is lg
Sponges from the Western Coast of
North America. Trans. roy. Soc, Can.,
vol. 124), 11S—138, Pl. 2-4.
Sponges from the Atlantic Coast of Can-
ada. Trans. roy. Soc. Can., (2) vol. 2 (4),
Po1—2 1 Pl 3.
1900a. Sponges from the coasts of northeastern
Canada and Greenland. Trans, roy. Soc.
Gan., (2) vol. 6 (4), 19-48, PI. 1-6:
1900b. Catalogue of the recent marine sponges
of Canada and Alaska. Ottawa Nat., vol.
14 (9), 153-172.

1893.

1894.

1896.

MARINE SPONGES

LAMOUROUX, J. V. F.

1824. Histoire naturelle des zoophytes, ou ani-
maux rayonnés. Jn: Encyclopédie méth-
odique, Vol. 2, Paris, Mme. Veuve
Agasse. 819p.

Lewy, JOSEPH

1857. (Note on Cliona.) Proc. Acad. nat. Sci.
Philad., 1856, vol. 8, 162-163.
1890. The boring-sponge, Cliona. Proc. Acad.

nat. Sci. Philad., 1889, 70-75.

LENDENFELD, ROBERT VON

1888. Descriptive catalogue of the sponges in
the Australian Museum, Sydney. Lon-
don, Taylor and Francis, 260p., 12 pl.

1898. Die Clavulina der Adria. Nova Acta

Leop. Carol., 1896, vol. 69 (1), 1-251,
Lelie Veils

LEVEAUX, MARCELLE

1939. La formation des gemmules chez les
Spongillidae. Ann. Soc. zool. Belg. vol.
70, 53-96, 21 figs.

LEvi, CLAUDE

1953a. Halisarca metschnikovi, n. sp., éponge
sans squelette des cdtes de France: ses
caractéres embryologiques. (Note pré-
liminaire.) Arch. Zool. exp. gén., vol.
90, Notes et revue (2): 87-91, 2 fig.
1953b. Déscription de Plakortis nigra nov. sp.
et remarqueés sur les Plakinidae (Démo-
sponges). Bull. Mus. Hist. nat. Paris (2)
vol. 25, 320-328, 4 fig.
Etude des Halisarca de Roscoff. Em-
bryologie et systématique des Démo-
sponges. Arch. Zool. exp. gén. vol. 93,
1-181, 62 fig.

LEVINSEN, G. M. R.
1893. Annulata, Hydroidae, Anthozoa, Porif-
era. In: Det videnskabelige udbytte af

Kanonbaaden “Hauchs” togter, 1883-
1886, p. 321-425, Pl. 1-3.

LIEBERKUHN, N.

1859. Neue Beitrage zur Anatomie der Spon-
gien. Arch. Anat. Physiol., Lpz., 1859,
515-529, Ll o-e

LINNE, CARL VON

1767. Systema Naturae. Holmiae, Laur, Sal-
vii. 12th ed., vol. 1 (2), 533-1327.

Luckt, BALDWIN AND Morton McCuTCHEON

1932. The living cell as an osmotic system
and its permeability to water. Physiol.
Rev., vol. 12, 68-139, 5 fig.

LUNDBECK, WILL

1902. Porifera. (Part I). Homorrhaphidae and
Heterorrhaphidae. Dan. Ingolf-Exped.,

vol. 6A (1), 104p.,.19 pl.

REFERENCES CITED

1905. Porifera. (Part II). Desmacidonidae
(Pars). Dan. Ingolf-Exped., vol. 6A (2),
219p., 7 fig., 20 pl.

Lunz, G. R., JR.

1935. A preliminary report of the survey of
the coastal waters of South Carolina.
Oyster Pest Control Investigations, U. S.
Bur. Fish., Prelim. Rep. (Mimeo-
graphed), 5p.

Maas, OTTo

1892. Die Metamorphose von Esperia lorenzi
O.S. nebst Beobachtungen an andern
Schwammlarven. Mitt. zool. Sta. Neapel,
vol. 10, 408-440, Pl. 27, 28.

McDoucaL.., K. D.

1943. Sessile marine invertebrates of Beaufort,
N. C. Ecol. Monogr., vol. 13 (3), 321-
374, 19 fig.

Maerz, A. J. AND M. R. PAUL

1950. A dictionary of color. 2d ed. New York,
McGraw-Hill, 208p., 56 pl.

MANTELL, GIDEON

1822. ‘The fossils of the South Down; or illus-

trations of the geology of Sussex. Lon-
don, Lupton Relfe, 327p., 42 pl.

Marsitul, L. F.

1725. Histoire physique de la mer. Amster-
dam, Boerhaave. 216p.

Mayr, Ernst, E. G. LINsLEy, AND R. L. USINGER

1953. Methods and principles of systematic
zoology. New York, McGraw-Hill. 328p.,
45 fig.

MENZEL, R. W.

1956. Annotated check-list of the marine
fauna and flora of the St. George’s
Sound—Apalachee Bay Region, Florida
Gulf Coast. Oceanogr. Inst. Fla, St. Univ.
Contr. No. 61, 78p.

MERCATI, MICHAELIS
I
1717. Metallotheca Vaticana. Rome, Jo: Ma-
riae Salvioni.
MEREJKOWSKY, C.

1879. Etudes sur les éponges de la Mer
Blanche. Mém. Acad. Sci. St.-Pétersb.,
(7) vol. 26 (7), 51p., 3 pl.

MIYAWAKI, MITSUHARU

1951. Notes on the effect of low salinity on
an actinian, Diadumene luciae, J. Fac.
Sci. Hokkaido Univ. (6) vol. 10 (2), 123-
126.

Moore, H. F.

1899. Report on the oyster-beds of Louisiana.
Rep. U. S. Comm. Fish., 1898, 49-100,
1 chart.

147

MULLER, KARL

1914. Gemmula-Studien und allgemein-biolog-
ische Untersuchungen an Ficulina ficus
Linné, Wiss. Meeresuntersuch., Abt.

Kiel, (2) vol. 16, 287-313, 10 fig., Pl. 4-7.
MULLER, O. F.

1776. Zoologiae danicae prodromus, seu ani-
malium Daniae et Norvegiae indige-
narum. Havniae, Typis Hallageriis.
282 p.

Narpo, G. D.

1833. Spongiariorum classification. Isis von
Oken, 1883, 519-523.

Rischiarimenti e rettificazioni ai generi
ed a qualche specie della famiglia de’
zoofitari sarcinoidi od alcionari stabilita
dal Sig. de Blainville. Nuovi Ann. Sci.
nat, R. C. Acad. Bologna (2) vol. 2, 460—
474.

Osservazioni sui generi di spongiali Sub-
urites e Litumena. Jn: Osservazioni post-
ume (Renier). (Reference not seen.)

1844,

1847.

Nasonovy, N. K.

1883. Zur Biologie und Anatomie der Clione.
Z. wiss Zool., vol. 39, 295-308, Pl. 18-19.
Sur l’éponge perforante Clione stationis
Nason. et le procédé du creusement des
galeries dans les valves des huitres. C. R.
Acad. Sci. Russ., 1924A, 113-115.

Les éponges perforantes de la famille
Clionidae de la Mer Noire et de la Mer
de Barentz. C. R. Acad. Sci. URSS.
1925A, 139-141.

NIKITIN, V. N.

1931. Die untere Planktongrenze und deren
Verteilung im Schwarzen Meer. Int. Rev.
Hydrobiol. vol. 25, 102-130, 4 fig.

1924.

1925.

1934. The Gudaut Oyster Bank. Trav. Sta.
pisc. biol. Géorgie vol. 1 (1), 51-179,
38 fig., 5 pl.

Novacek, R.

1930. Biologické faktory pri vzniku dolomitt.
Sborn. prir. ¢es. Akad., vol. 7, 53-69.

OLD, M. C.

1941. The taxonomy and distribution of the
boring sponges (Clionidae) along the
Atlantic Coast of North America. Publ.
Chesapeake biol. Lab., vol. 44, 30p., 13
pl., 1 map.

The boring sponges and their effect on
shellfish culture. Ann. Meeting nat.
Shellfisheries Ass. Philad. 3p. (Mimeo-
graphed).

1942.

Outvi, A. GIUSEPPE

1792. Zoologia adriatica ossia catalogo ragion-
ato degli animali del Golfo e delle La-
gune di Venezia; preceduto da una dis-

148

sertazione sulla storia fisica e naturale
del Golfo; e accompagnato da memorie,
ed osservazioni di fisica storia naturale
ed economia. Bassano. 334p., 9 pl.

Orton, J. H.

1937. Oyster biology and oyster-culture. Lon-
don, Edward Arnold and Co. 211p., 57
fig.

PALLAs, P. S.

1766. Elenchus zoophytorum sistens generum
adumbrationes generaliores et specierum
cognitarum succinctas descriptiones cum
selectis auctorum synonymis. Hagae-
Comitum, Petrum van Cleef. 451p.

PALMHERT, H. W.

1933. Beitrage zum Problem der Osmoregula-
tion einiger Hydroidpolypen. Zool. Jb.
Abt. allg. Zool. Physiol., vol. 53, 212-
260, 18 fig.

PANTIN, C. F. A.
1954. The recognition of species. Sci. Progr.
vol. 168, 587-598.

Parker, G. H.

1910. The reactions of sponges, with a con-
sideration of the origin of the nervous
system. J. exp. Zool., vol. 8 (1), 1-41, 3
fig.

Parr, A. iE.

1933. A geographic-ecological analysis of the
seasonal changes in temperature condi-
tions in shallow water along the At-
lantic Coast of the United States. Bull.
Bingham oceanogr. Coll., vol. 4 (3), 90p.,
28 fig.

PEARL, RAYMOND AND J. R. MINER

1935. Experimental studies on the duration
of life. XIV. The comparative mortality
of certain lower organisms. Quart. Rev.
Biol., vol. 10, 60-79, 5 fig.

PEARSE, A. S. AND G. W. WHARTON

1938. The oyster “leech,” Stylochus inimicus
Palombi, associated with oysters on the
coasts of Florida. Ecol. Monogr., vol. 8
(4), 605-655, 37 fig.

Poret, J. L. M.

1789. Reise in die Barbarey oder Briefe aus
Alt-Numidien geschrieben in den Jah-
ren 1785 und 1786. . . . Nebst einem
Versuche tiber die Naturgeschichte di-
eses Landes. Zweiter Theil. Strassburg,
Akademische Buchhandlung, 267p., 4 pl.

POMERAT, C. M. AND E. R. REINER

1942. ‘The influence of surface angle of light
on the attachment of barnacles and
other sedentary organisms. Biol. Bull.
Woods Hole, vol. 82, 14-25, 3 fig.

MARINE SPONGES

POuURBAIX, NELLY

1935. Formation histochimique des gemmules
d’éponges. Ann. Soc. zool. Belg. vol. 66,
Somos

PROCTER, WILLIAM

1933. Porifera. In: Biol. Surv. Mt. Desert Re-
gion. Part V. Marine Fauna. Philad.
Wistar Inst., p. 78-115.

RAFINESQUE, C. S.

1819. Descriptions of species of sponges ob-
served on the shores of Long Island.
Amer. J. Sci., vol. 1 (11), 149-151.

Ray, JOHN

1724. Synopsis methodica stirpium britanni-
carum: Tum indigenis, tum in agris
cultis locis suis dispositis; additis gen-
erum characteristicis, specierum descrip-
tionibus et virium epitome. London,
Gulielmus & Joannis Innys. 482p., 24

pl.
REDEKE, H. C.

1922. Zur Biologie der niederlandischen Brack-
wassertypen. Bijdr. Dierk., vol. 22, 329-
335.

REMANE, ADOLF

1934. Die Brackwasserfauna (mit besonderer
Berticksichtigung der Ostsee). Zool. Anz.
Suppl. vol. 7, 34-74, 4 fig.

RENOvuF, Loutis

1936. The importance of field notes to the
study of Porifera. C, R. XIII® Congr. in-
ternat. Zool. Lisbonne, 1935, 831-840,
2 fig.

Rinrey, G. A.

1948. Hydrography of the Western Atlantic;
the Long Island and Block Island
Sounds. Tech. Rep. Woods Hole
oceanogr. Instn, vol. 11, 30p., 32 fig.

1956. Oceanography of Long Island Sound,
1952-1954. II. Physical Oceanography.
Bull. Bingham oceanogr. Coll. vol. 15,
15-46, 14 fig.

ROCHE, JEAN AND A. TIXIER-DURIVAULT

1951. Rapports des Gérardiidés avec les Zoan-
thides et les Antipathaires. Bull. Mus.

Hist. nat. Paris (2) vol. 23, 402-409, 1 fig.

SCHMIDT, E. O.

1866. Zweites Supplement der Spongien des
adriatischen Meeres. Enthaltend die
Vergleichung der adriatischen und _ bri-
tischen Spongiengattungen. Leipzig, Wil-
helm Engelmann. 23p., 1 pl.
Grundziige einer Spongien-Fauna des
atlantischen Gebietes. Leipzig; Wilhelm
Engelmann. 88p., 6 pl.

1870.

REFERENCES CITED

SCHMIDTLEIN, R.

1878. Beobachtungen tiber Trachtigkeits- und
Eiablage-Perioden verschiedener Seeth-
iere. Mitt. zool. Sta. Neapel, vol. 1, 124-
136.

SCHULZ, BRUNO

1932. Einfiihrung in die Hydrographie der
Nord- und Ostsee. Tierwelt N.-u. Ostsee,
Lief. 21, Teil Id,, 45-88, 25 fig.

SEURAT, L. G.

1934. Formations littorales et estuaires de la
Syrte Mineure. (Golfe de Gabés). Bull.
Sta. océanogr. Salammbd, no. 32, 65p.,
7 fig., 1 chart.

Smi1TH, R. I., e¢ al.

1954. Intertidal invertebrates of the Central
California Coast. Berkeley. University of
California Press. 446p., 138 fig.

SmiTH, S. I., AND O. HARGER

1876. Report on the dredgings in the region
of St. George’s Banks, in 1872. Trans.
Conn. Acad. Arts Sci., vol. 3 (1), 1-57,
Pi. 1-8.

SuMNER, F. B., R. C. OsBuRN, AND L. J. COLE

1913. A biological survey of the waters of
Woods Hole and vicinity. III. A cata-
logue of the marine fauna of Woods
Hole and vicinity. Bull. U. S. Bur. Fish.,
1911, vol. 31 (2), 545-860.

SvERDRUP, H. U., M. W. JOHNSON, AND
R. H. FLEMING

1946. The oceans: Their physics, chemistry,
and general biology. New York, Pren-
tice-Hall, Inc., 1087p., 265 fig., 7 charts.

SWARTSCHEWSKY, B.

1905. Beitrag zur Kenntnis der Schwamm-
Fauna des Schwarzen Meeres. Mém.
Soc. Nat. Kiew., vol. 20 (1), 1-59, PI. 1-7.

1906. Beitrag zur Spongien-Fauna des Weissen
Meeres. Mém. Soc. Nat. Kiew, vol. 20
(2), 307-371, Pl. 10-16.

THIELE, JOHANNES

1898. Studien tiber pazifische Spongien. Zoolo-
gica, Stuttgart, vol. 24 (1), 72p., 8 pl.

1905. Die Kiesel- und Hornschwéamme der
Sammlung Plate. Zool. Jb., Suppl. vol. 6,
407-496, pl. 27-33.

THOMSON, J. A.

1887. On the structure of Suberites domun-
cula Olivi (O. S.), together with a note
on peculiar capsules found on the sur-
face of Spongelia. Trans. roy. Soc.

Edinb., vol. 33, 241-245, Pl. 16-17.

149

TorsENT, EMILE
1887. Contribution 4 l’étude des Clionides.
Arch. Zool. exp. gén., (2) vol. 5 Bis, 1-
166, 7 pl.
Quelques Spongiaires du Banc de Cam-
péche et de la Pointe-a-Pitre. Mem. Soc.
zool. Fr., vol. 2, 30-52, 12 fig.
Diagnoses d’éponges nouvelles de la
Méditerranée et plus particuli¢rement
de Banyuls. Arch. Zool. exp. gén., (2)
vol. 10, Notes et revue (6): xvii-xxviii.
Application de la taxonomie actuelle a
une collection de spongiaires du Banc
de Campéche et de la Guadeloupe dé-
crite précédemment. Mém. Soc. zool. Fr.,
vol. 7, 27-36.
Etude sur la faune des Spongiaires du
Pas-de-Calais suivie d’une application
de la nomenclature actuelle 4 la mono-
graphie de Bowerbank. Rev. Biol. du
Nord de la France, vol. 7, 6-28.
Spongiaires de la Baie d’Amboine. Rev.
suisse Zool. vol. 4 (3), 421-487, Pl. 18-21.
Etude monographique des Spongiaires
de France. IIT. Monaxonida (Hadrome-
rina). Arch. Zool. exp. gén., (3) vol. 8,
1-331, Pl. 1-8:
Sur les affinités des Halichondria et la
classification des Halichondrines d’aprés
leurs formes larvaires. Arch. Zool. exp.
gén., (5) vol. 7. Notes et revue (1), i-xv,
4 fig.
1925a. Etudes de Spongiaires du Golfe de
Naples. Arch. Zool. exp. gén. vol. 63,
623-725, 27 fig., Pl. 8.
1925b. Eponges de l’Etang de Thau. Bull. Inst.
océanogr. Monaco, no. 452, 19p., 2 fig.
1932. Notes sur les Clionides. Arch. Zool. exp.
gén., vol. 74, 549-579, 11 fig.
Eponges de Lamarck conservées au Mu-
séum de Paris. Fin. Arch. Mus. Hist.
nat. Paris, (6) vol. 10, 1-60, 5 fig., 3 pl.

TuzET, ODETTE

1889.

1892.

1894.

1895.

1897.

1900.

1911.

1933.

1930. Sur la fécondation de l’éponge siliceuse,
Cliona viridis Schmidt. C. R. Acad. Sci.,
Paris. vol. 191, 1095-1097.

1948. Les premiers stades du développement
de Leucosolenia botryoides Ellis et So-
lander et de Clathrina (Leucosolenia)
coriacea Mont. Ann. Sci. nat. Zool. (11)
vol. 10, 103-114.

VALIKANGAS, ILMARI

1933. Uber die Biologie der Ostsee als Brack-
wassergebiet. Verh. int. Ver. Limnol.,
vol. 6, 62-112, 6 fig., 1 chart.

VANEY, CLEMENT AND A. ALLEMAND- MARTIN

1918. Contribution a l’étude de la larve de
V’Hippospongia equina des cétes de Tu-
nisie. C. R. Acad. Sci., Paris. vol. 166,
82-84.

150

VERRILL, A. E.

1871. Brief contributions to zoology from the
Museum of Yale College. No. XVI. On
the distribution of marine animals on
the southern coast of New England.
Amer. J. Sci., (3) vol. 2, 357-362.

1874a. Brief contributions to zoology from the
Museum of Yale College. No. XXVI. Re-
sults of recent dredging expeditions on
the coast of New England. No. 4.
Amer. J. Sci., (3) vol. 7, 38-46.

1874b. Explorations of Casco Bay by the U. S.

Fish Commission, in 1873. Proc. Amer.

Ass. Adv. Sci., vol. 22 (2), 340-395, 6 pl.

Brief contributions to zoology from the

Museum of Yale College. No. XXXIII.

Results of dredging expeditions off the

New England Coast in 1874. Amer. J.

Set(3) Voll0N30-—43, 081) 34.

Note on borings of a sponge in Italian

marble. Amer. J. Sci., (3) vol. 16, 406.

List of marine Invertebrata from the

New England Coast, distributed by the

U. S. Commission of Fish and Fisheries.

Proc. U. S. nat. Mus., 1879, vol. 2, 227-

Boar

VeRRILL, A. E. AND S. I. SMITH

1873. Report upon the invertebrate animals
of Vineyard Sound and adjacent waters,
with an account of the physical char-

acters of the region. Rep. U. S. Comm.
Fish., 1871-1872, 295-778, 4 fig., 38 pl.

VinoGRADOV, A. P.

1953. The elementary chemical composition of
marine organisms. Mem. Sears Fdn Mar.
Res., vol. 2, 647p.

VOLZ, PETER

1939. Die Bohrschwamme (Clioniden) der

Adria. Thalassia, vol. 3 (2), 1-64, 16 fig.,
5 pl., 3 maps.

1875.

1878.

1880.

MARINE SPONGES

VosmaeEr, G, C. J.

1882. Report on the sponges dredged up in
the Arctic Sea by the “Willem Barents”
in the years 1878 and 1879. Niederl.
Arch. Zool. Suppl. vol. 1, 1-58, Pl. 1-4.
The Porifera of the Siboga-Expedition.
II. The Genus Spirastrella. Siboga-Ex-
ped., vol. 6A, 69p., 14 pl.

The sponges of the Bay of Naples: Pori-
fera Incalcaria. Ed. by C. S. Vosmaer-
Roéll and M. Burton. The Hague: Mar-
tinus Nijhoff. 1933-1935. 3 vols., 828p.,
69 pl.

191T

1933.

WELLs, G. P. AND I. C. LEDINGHAM

1940. Physiological effects of a hypotonic en-
vironment. I, The action of hypotonic
salines on isolated rhythmic prepara-
tions from polychaete worms (Arenicola
marina, Nereis diversicolor, Perinereis
cultrifera). J. exp. Biol., vol. 17, 337-352,
8 fig.

WHITEAVES, J. F.

1901. Catalogue of the marine Invertebrata of
eastern Canada. Geol. Surv. Can., Ot-
tawa, 271p.

Witson, H. V.

1894. Observations on the gemmule and egg
development of marine sponges. J.
Morph., vol. 9 (3), 277-406, Pl. 14-25.

1912. Development of sponges from dissoci-
ated tissue cells. Bull. U. S. Bur. Fish.,
1910, vol. 30, 1-30, 5 pl.

1935. Some critical points in the metamorpho-

sis of the Halichondrine sponge larva.
J. Morph., vol. 58, 285-345, 4 pl.

ZEUTHEN, ERIK

1939. On the hibernation of Spongilla lacu-
stris (L.). Z. vergl. Physiol., vol. 26, 537—
547, 3 fig.

SYSTEMATIC INDEX

Numbers in italics indicate page on which subject is illustrated; plate references are in capital

Roman numerals.

Adocia tubifera, 71, 72, 78, 99

Aka trachys, 132

albescens, Spongia, 1

albicans, Cliona, 137

Alcyonio minore in forma di fico frutto, 15

Alcyonium, compactum, 4; domuncula, 15;
ficus, 4, 13, 14, 15; quintam antiquorum,
14; tuberosum forma ficus, 14

Anchinoe fictitia, 121

anomala, Isodictya, 45

Anthosigmella, 13

arbuscula, Chalina, 2, 52, 58, 59, 62; Haliclona,
52, 56

arctica, Rinalda, 94

Ascortis fragilis, 2

birotulata, Iotrochota, 107
bowerbanki, Halichondria, x, 3, 24-36, 27, 35,
80, 87, 102-104, 103, 105, II, III, IV

canaliculata, Haliclona, 3, 73-76, 74, 77, V

caribboea, Cliona, 16, 17

carnosus, Suberites, 13, 14

carolinensis, Lissodendoryx, 41

celata, Cliona, 2, 3, 16-18, 19, 20, 79, 87-93, 104,
106-113, 114-119, 123-127, 126, 129-141,
USK, WEY hs

cespitosa, Spongia, 1, 62

Chalina, arbuscula, 2, 52, 58, 59, 62; ficus, 4;
flemingii, 71, 72; montaguii, 71, 72; oculata,
2, 62

Choanites, 13; ficus, 16

Choanitidae, 13

ciliata, Grantia, 2

cinerea, Reniera, ix

cladonia, Spongia, 1

Clathria delicata, 37, 41

Clathrina, x

Cliona, albicans, 137, caribboea, 16, 17; celata,
2, 3, 16-18, 19, 20, 79, 87-93, 104, 106-113,
114-119, 123-127, 126, 129-141, 136, 137, I;
euryphylla, 132; lobata, 2, 3, 16, 19-21, 19,
127, 129, 1305 1325 133,134; pontica; 130;
robusta, 129, 131; rovignensis, 137; schmidti,
16, 132, 137; spirilla, 23, 129, 131; stationis,
80, 89, 138; sulphurea, 2, 17; truitti, 2, 3,
22-24, 125-129, 126, 130, 131; vastifica, x, 2,
3, 19, 21, 21-23, 79, 87, 93-94, 104, 106-109,
120, 124-127, 126, 129-141, 136, 137; vermi-
fera, 137, 138; viridis, 16, 80, 90, 137, 138

Clionidae, 3, 16

Cliothosa hancocki, 80, 137

coalita, Halichondria, 24, 36; Spongia, 36

compacta, Halichondria, 4, 15; Suberites, 4, 13,
16

compactum, Alcyonium, 4

compressa, Grantia, 79, 121

concinnus, Suberites, 7

coriacea, Leucosolenia, 121

deichmannae, Isodictya, 3, 45-52, 47, 48, V, VI
deichmanni, Neosperiopsis, 45

delicata, Clathria, 37, 41

Dendroceratida, 3

Desmacidonidae, 3, 44

domuncula, Alcyonium, 15

domunculus, Suberites, 5-7, 9-16, 10, 37
dujardini, Halisarca, 121

dura, Reniera, var. ficiformis, 14

Ephydatia, 121

Esperiopsis, 45; laxa, 46, 48; quatsinoensis, 45,
46, 48; rigida, 46, 48; vancouverensis, 46

euryphylla, Cliona, 132

farinaria, Halichondria, 4; Suberites, 4

farinarius, Suberites, 4

ficiformis, Petrosia,
Spongia, 4, 14

fictitia, Anchinoe, 121; Microciona, 121

Ficulina, 13; ficus, 4, 14, 16

ficus, Alcyonium, 4, 13, 14, 15; Chalina, 4;
Choanites, 16; Ficulina, 4, 14, 16; Hali-
chondria, 4, 15; Hymeniacidon, 4, 15;
Suberites, 1, 3-16, 10, I

ficus?, Spongia, 15

flemingii, Chalina, 71, 72

fragilis, Ascortis, 2

14; Reniera dura, 14;

Gellius, 56

Grantia, ciliata, 2; compressa, 79, 121
griffithsii, Raphyrus, 91

grossa, Halichondria, 36

Hadromerina, 3

Halichondria, ix, x, 68; bowerbanki, x, 24-36,
27, 35, 80, 87, 102-104, 103, 105, II, III, IV;
coalita, 24, 36; compacta, 4, 15; farinaria,
4; ficus, 4, 15; grossa, 36; isodictyalis, 41;
luxurians, 36; montaguii, 71; panicea, ix, x,
2, 24, 29, 30-34, 35, 36, 80, 104, II; suberea,
4, 11, 15; virgultosa, 4, 15

Halichondriidae, 3, 24

Halichondrina, 3, 24

152

Haliclona, ix, x, 37; arbuscula, 52, 56; canalicu-
lata, 3, 73-76, 74, 77, V; implexa, 59;
loosanoffi, 2, 3, 62-72, 64-66, 87, 94-102,
98-101, 103, 104-105, XI, XII; montagui,
59; oculata, ix, x, 1, 3, 36, 52-62, 60, 61,
VII, VIII, IX, X; oculata tavaresi, 59;
palmata, 58, 71, 72; permollis, ix, x, 68-71,
[Pe

Haliclonidae, ix, 3, 52

Halina suberea, 4

Halisarca, x, 68; dujardini, 121; sp.?, 2, 3

Halisarcidae, 3

hancocki, Cliothosa, 80, 137

Haplosclerina, 3, 44

heliophila, Hymeniacidon, 122; Stylotella, 122

heterofibrosa, Reniera, 71, 72

Homoeodictya, 45

Hymeniacidon, ficus, 4, 15; heliophila, 122;
suberea, 7; subereum, 4, 11; virgultosa, 4, 15

implexa, Haliclona, 59

infundibuliformis, Isodictya, 45

Iotrochota birotulata, 107

Isodictya, 44-45; anomala, 45; deichmannae,
3, 45-52, 47, 48, V, VI; infundibuliformis,
45; lobata, 45; normani, 45; (palmata?), 2;
palmata, 45, 46, 47, 49, 51, 52, VI

isodictyalis, Halichondria, 41; Lissodendoryx, 3,
41-44, 42, IV

lacustris, Spongilla, 76

latus, Suberites, 4, 11, 16

laxa, Esperiopsis, 46, 48

leptoderma, Lissodendoryx, 41; Tedania, 41

Leucetta losangelensis, 78

Leucosolenia, x, 121; coriacea, 121

Lissodendoryx, carolinensis, 41; isodictyalis, 3,
41-44, 42, IV; leptoderma, 41

lobata, Cliona, 2, 3, 16, 19-21, 19, 127, 129, 130,
132, 133, 134; Isodictya, 45

loosanoffi, Haliclona, 2, 3, 62-72, 64-66, 87, 94—
102, 98-101, 103, 104-105, XI, XII

losangelensis, Leucetta, 78

liitkeni, Suberites, 4, 13

luxurians, Halichondria, 36

mammillaris, Polymastia, 94

Microciona, fictitia, 121; prolifera, 1, 2, 3, 36-41,
40, 78, 87, 104, 107, IV

Microcionidae, 3, 36

mollis, Reniera, 71, 72; Thoosa, 80, 137

montaguii, Chalina, 71, 72; Halichondria, 71;
Haliclona, 59

montalbidus, Suberites, 4, 16

montiniger, Suberites, 4, 7

Mycale, 56

Myxilla, 56

Myxillidae, 3, 41

Neosperiopsis deichmanni, 45
normani, Isodictya, 45
nuttingi, Rhabdodermella, 78

MARINE SPONGES

oculata, Chalina, 2, 62; Haliclona, ix, x, 1, 3,
36, 52-62, 60, 61, VII, VIII, IX, X; Hali-
clona, tavaresi, 59

ostracina, Spongia, 1, 37, 41

(palmata?), Isodictya, 2

palmata, Haliclona, 58, 71, 72; Isodictya, 45, 46,
47, 49, 51, 52, VI; Spongia, 58

panicea, Halichondria, ix, x, 2, 24, 29, 30-34,
35, 36, 80, 104, II

Papillina suberea, 91

permollis, Haliclona, ix, x, 68-71, 72

Petrosia ficiformis, 14

placenta, Suberites, 4, 13

plana, Reniera, 76

Poecilosclerina, 3, 36

Polymastia, 56; mammillaris, 94

pontica, Cliona, 130

prolifera, Microciona, 1, 2, 3, 36-41, 40, 78, 87,
104, 107, IV; Spongia, 36, 37, 41

pulmonaria, Synoicum, 13

purpurea, Spirastrella, 93

quatsinoensis, Esperiopsis, 45, 46, 48
quintam antiquorum, Alcyonium, 14

raphanus, Sycon, 79

Raphyrus griffithsii, 91

Reniera, cinerea, ix; dura var. ficiformis, 14;
heterofibrosa, 71, 72; mollis, 71, 72; plana,
76; tubifera, 71, 72, 78

Rhabdodermella nuttingi, 78

rigida, Esperiopsis, 46, 48

Rinalda arctica, 94

robusta, Cliona, 129, 131

rovignensis, Cliona, 137

schmidti, Cliona, 16, 132, 137

Spheciospongia, 13

Spirastrella, 93, 132, 139; purpurea, 93

Spirastrellidae, 13

spirilla, Cliona, 23, 129, 131

Spongia, albescens, 1; cespitosa, 1, 62; cladonia,
1; coalita, 36; ficiformis, 4, 14; ficus?, 15;
ostracina, 1, 37, 41; palmata, 58; prolifera,
36, 37, 41; suberosa, 36; sulphurea, 17;
urceolata, 37, 41; virgata, 1; virgultosa, 15

Spongilla, 37, 63, 121; lacustris, 76

stationis, Cliona, 80, 89, 138

Stylotella heliophila, 122

suberea, Halichondria, 4, 11, 15; Halina, 4;
Hymeniacidon, 7; Papillina, 91; Suberites,
4, 13, 16

subereum, Hymeniacidon, 4, 11

Suberites, 15; carnosus, 13, 14; compacta, 4, 13,
16; concinnus, 7; domunculus, 5-7, 9-16,
10, 37; farinaria, 4; farinarius, 4; ficus, 1,
3-16, 10, I; latus, 4, 11, 16; liitkeni, 4, 13;
montalbidus, 4, 16; montiniger, 4, 7; pla-
centa, 4, 13; suberea, 4, 13, 16; typus, 15;
virgultosa, 4; volubilis, 15

SUBJECT INDEX

Suberitidae, 3, 13

suberosa, Spongia, 36

Suburites, 15

sulphurea, Cliona, 2, 17; Spongia, 17
Sycon raphanus, 79

Synoicum pulmonaria, 13

tavaresi, Haliclona oculata, 59

Tedania leptoderma, 41

Thoosa mollis, 80, 137

trachys, Aka, 132

truitti, Cliona, 2, 3, 22-24, 125-129, 126, 130,
131

tuberosum forma ficus, Alcyonium, 14

tubifera, Adocia, 71, 72, 78, 99; Reniera, 71, 72,
78

153
typus, Suberites, 15

urceolata, Spongia, 37, 41

vancouverensis, Esperiopsis, 46

vastifica, Cliona, x, 2, 3, 19, 21, 21-23, 79, 87,
93-94, 104, 106-109, 120, 124-127, 126, 129-
141, 136, 137; Vioa, 21

vermifera, Cliona, 137, 138

Vioa vastifica, 21

virgata, Spongia, |

virgultosa, Halichondria, 4, 15; Hymeniacidon,
4, 15; Spongia, 15; Suberites, 4

viridis, Cliona, 16, 80, 90, 137, 138

volubilis, Suberites, 15

SUBJECT INDEX

Numbers in italics indicate page on which subject is illustrated; plate references are in capital

Roman numerals.

Amphipods, living in Suberites ficus, 5

Anthozoa, biochemical characteristics and clas-
sification, x

Asexual reproduction, Cliona vastifica, 93-94;
Polymastia mammillaris, 94

Biochemical characteristics, value in system-
atics, x

Boring process of clionids, 89

Breeding seasons of sponges, 78-85

Calcareous algae, association with, Cliona ce-
lata, 90; C. viridis, 90

Cell cords, Haliclona canaliculata, 73; H. loosa-
noffi, 63, 64; H. oculata, 60; Microciona
prolifera, 37; spongillids, 37, 63; Suberites
domunculus, 13, 37

Chalina flemingii, compared with Haliclona
loosanoffi, 71, 72

Chalina montaguti, compared with Haliclona
loosanoffi, 71, 72

Choanocytes, loss of, in winter, ix, 30, 76

Cliona celata, distribution, 17, 77; distribution
compared with C. vastifica, 124-125, 126,
133, 135-140, 136; growth stages, 16, 90-93,
I; microscleres in young colonies, 17; papil-
lar structure, 107-108; reversion of free-
living stage to boring habit, 91-93; shell
perforations, size of, 17; spicule dimen-
sions, 18; synonymy, 16

Cliona lobata, distribution, 20-21, 77; shell
perforations, size of, 19; spicule dimen-
sions, 20; spicule structure, 19; synonymy,
19

Cliona robusta, spicule dimensions, 131

Cliona spirilla, spicule dimensions, 131

Cliona stationis, spicule dimensions, 131

Cliona truitti, distribution, 24, 77; shell per-
forations, size of, 24; spicule dimensions,
23, 131; spicule structure, 22

Cliona vastifica, distribution, 22, 77; distribu-
tion compared with C. celata, 124-125, 126,
133, 135-140, 136; papillar structure, 108;
spicule dimensions, 23, 131; spicule struc-
ture, 27, 22; synonymy, 21

Clionids, boring process of, 89; control of, 140—
141; occurrence in different substrata, 132;
origin of boring habit, 93, 139; salinity in-
dicators, usefulness as, 125; vertical distri-
bution at Rovigno, 137, 138

Contractile vacuoles, in marine sponges, 121; in
spongillids, 121, 122

Current, water, influence on larval settling in
Adocia tubifera, 78

Depth, influence on larval settling in Haliclona
loosanoffi, 99, 100

Distribution, dependence on length of growing
season in Haliclona loosanoffi, 102

Eggs, Cliona celata, 17; Halichondria bower-
banki, 30; Haliclona loosanoffi, 66-67; H.
oculata, 59

Embryos, Haliclona loosanoffi, 66-67; H. ocu-
lata, 59

Esperiopsis laxa, compared with Isodictya
deichmannae, 46, 48

154

Esperiopsis quatsinoensis, compared with Iso-
dictya deichmannae, 46, 48; spicule dimen-
sions, 51

Esperiopsis rigida, compared with JIsodictya
deichmannae, 46, 48

Esperiopsis vancouverensis, compared with Iso-
dictya deichmannae, 46, 48

Gemmules, Cliona vastifica, 140; Haliclona
loosanoffi, 63, 65, 66, 97-98, 101, 102, XII;
Suberites domunculus, 5; S. ficus, 5

Growth, Adocia tubifera, 78; Haliclona loosa-
noffi, 94-96; Leucetia losangelensis, 78;
Microciona prolifera, 78; Rhabdodermella
nuttingi, 78

Halichondria bowerbanki, colony form, III, IV;
comparison with, H. grossa, 36, H. luxuri-
ans, 36, H. panicea, 30-34, 35, II; dermal
skeleton, 26-27, 30, 31, 33-34, 35, II; dis-
tribution, 30, 77; intertidal distribution,
104; spicule dimensions, 28, 31; spicules of
American and European populations com-
pared, 26-27, 30; spicule structure, 26, 27;
synonymy, 36; winter degeneration, 30;
winter killing, 102, 104

Halichondria grossa, compared with H. bower-
banki, 36

Halichondria luxurians, compared with H.
bowerbanki, 36

Halichondria montaguii, compared with Hali-
clona loosanoffi, 71

Halichondria panicea, comparison with H.
bowerbanki, 30-34, 35, II; dermal skeleton,
33-34, 35, II; spicule dimensions, 29, 31

Haliclona canaliculata, colony form, V; com-
parison with, H. loosanoffi, 76, Reniera
plana, 76; distribution, 76, 77; skeletal
framework, 73, 74; spicule dimensions, 75;
spicule structure, 74, 76; winter degenera-
tion, 76

Haliclona loosanoffi, anatomy, 63, 64; colony
form, XI, XII; comparison with other
haliclonids, 68-72, 76; distribution, 67, 77;
life history summarized, 103; skeletal
framework, 63, 65; spicule dimensions, 69-—
70; spicule structure, 63, 65; survival data,
94, 97-98; variations in spicule size, 67-68;
winter killing, 99, 100, 101

Haliclona oculata, colony form, VII, VIII, IX,
X; distribution, 60, 62, 77; skeletal frame-
work, 59, 60, 61; spicule dimensions, 54-55;
synonymy, 52; variations in form, 52-53

Haliclona palmata, compared with H. loosa-
nope, MM, 12

Haliclona permollis, compared with H. loosa-
noffi, 68-71, 72

Hymeniacidon heliophila, reactions to lowered
salinity, 122, 123

Isodictya deichmannae, colony form, V, VI; dis-
tribution, 46, 52, 77; skeletal framework,

MARINE SPONGES

48; spicule dimensions, 50-51; spicule struc-
ture, 45-46, 47

Isodictya palmata, colony form, VI; distribu-
tion, 46, 49, 52; skeletal framework, 49;
spicule dimensions, 51; spicule structure,
46, 47

Larvae, of Halichondria bowerbanki, 27, 30,
32, 34, American and European popula-
tions compared, 32, 34, 36, comparison with
H. panicea, 32, 34; of Halichondria panicea,
32, 34, comparison with H. bowerbanki, 32,
34; interspecific differences, x

Larval settling period, Adocia tubifera, 78;
Cliona celata, 87-88; Cliona vastifica, 87,
93; Halichondria bowerbanki, 87, 102, 103;
Haliclona loosanoffi, 87, 94-95, 98-99; Mi-
crociona prolifera, 78, 87, 104

Life cycles, interspecific differences, ix

Life table for Haliclona loosanoffi, 94-97

Light, influence on larval settling, Adocia tubi-
fera, 79, 99; Halichondria bowerbanki, 98;
Haliclona loosanoffi, 98

Lissodendoryx isodictyalis, colony form, IV;
distribution, 44, 77; spicule dimensions,
43-44; spicule structure, 41, 42

Long Island Sound, salinity of, 106

Microciona prolifera, colony form, IV; distribu-
tion, 41, 77; spicule dimensions, 38-39;
spicule structure, 37, 40

Milford Harbor, salinity variations, 121

Osmoregulation, in coelenterates, 122; in
sponges, 121-123

Oysters, extent of infection by clionids in Long
Island Sound, 88, 133-135; preferred as
substratum by Cliona celata larvae, 88

Polyhaline zone in Long Island Sound, 106
Predation of sponges by molluscs, 97, 104

Reniera heterofibrosa, compared with Hali-
clona loosanoffi, 71, 72

Reniera mollis, compared with Haliclona loosa-
noffi, 71, 72

Reniera plana, compared with Haliclona cana-
liculata, 76

Salinity, correlation with spicule form, x, 129-
130; influence on coalescence of disso-
ciated cells, 107; influence on distribution,
Cliona celata, 124, 125, 126, 127, 129, 136,
Cliona lobata, 127, 129, Cliona robusta,
129, Cliona spirilla, 129, Cliona truitti, 125,
126, 127, 128, 129, Cliona vastifica, 125, 126,
127, 128-131, 136

Salinity tolerance, comparison between Cliona
celata and C. vastifica, 124; experiments
on, C. celata, 108-113, 114-119, 124, C.
vastifica, 109-110, 120, 124, methods of
study, 107, survival index, 108; mechanism
of, in Cliona, 122-123

SUBJECT INDEX

Silicon concentration, correlation with spicule
form, x, 76

Skeleton, intraspecific variation, ix, 59, 60, 61

Species, aesthetic recognition of, ix

Spicules, environmental variations, x; form of,
correlated with salinity, x, 129-130, corre-
lated with silicon concentration, x, 76, cor-
related with temperature, 49; size of, cor-
related with temperature, x, 40, 52-54, 56;
seasonal variation in size, 30, 76

Sponges, as “plants,” 1

Spongillids, cell cords, 37, 63; contractile
vacuoles, 121, 122; spicule form corre-
lated with silicon concentration, x, 76

Spongin, chemical composition and classifica-
tion, x; variation in amount, in Haliclona
oculata, 59, 60, 61, in Reniera cinerea, ix

Sterols and sponge classification, x, 13

155

Suberites domunculus, compared with S. ficus,
5-13; distribution, 12; nomenclature, 15—
16; spicule dimensions, 9; spicule struc-
ture, 5, 7, 10; spongin fibers, 6

Suberites ficus, colony form, I; compared with
S. domunculus, 5-13; distribution, 12, 16,
77; nomenclature, 13-16; relationships, 13;
spicule dimensions, 8; spicule structure, 7,
10, 11; spongin fibers, 6; synonymy, 4

Substratum, color of, influence on larval settling
of Haliclona loosanoffi, 99

Temperature, water, correlation with larval
settling in Haliclona loosanoffi, 101; corre-
lation with spicule form, 49; correlation
with spicule size, x, 40, 52-54, 56

Winter degeneration of sponges, ix, 30, 37, 76
Winter killing of sponges, 99-101, 102, 104

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REA a

. Cliona celata. Living colony, a-stage, growing in shell of Crassostrea vir-

ginica. New Haven Harbor; 7 meters. Photographed in running sea
water in laboratory. x 7/10.

. Portion of same colony showing incurrent papillae and oscule. x 2.8.

. Cliona celata. f-stage, overgrowing shell of Crassostrea virginica. New

Haven Harbor; 7 meters. x 3/4.

. Cliona celata. y-stage, photographed soon after removal from water. New

Haven Harbor, 9 meters. x 1/4.

. Suberites ficus. Block Island Sound; 40 meters. YPM +795 (dried speci

men). x 1/3.

. Cultch bag used in studies of sponge settling. x 1/8.

PEATE 2
Fig. 1. Halichondria panicea. Pattern of dermal skeletal tracts. Plymouth, Eng-
land, YPM 322097..x 24.
Fig. 2. Halichondria bowerbanki. Pattern of dermal skeletal tracts. Double Beach
(Branford), Conn. YPM 7£2019F. x 24.

oC
>

jl bye QI Ea 3
Variations of colony form in Halichondria bowerbanki
(All photographs from specimens preserved in alcohol.)

Figs. 1, 2, 3. Colonies with massive base and both rounded and lamellate vertical
branches; viewed from the side. Figs. 1, 2: Lagoon Pond, Martha’s
Vineyard, YPM #940. x 1/2. Fig. 3: Double Beach (Branford), Conn.
YPM 222019D: x 1/2.

Fig. 4. Colony with massive base and rounded, anastomosing branches; viewed
from side. Hammonasset, Conn. YPM #2024D. x 1/2.

Fig. 5. Colony with massive base and anastomosing, rounded branches; viewed
from above. Hammonasset, Conn. YPM 7£2024E. x 1/2.

Figs. 7, 8. Colonies with vertical processes which do not anastomose. Double
Beach (Branford), Conn. YPM #2019E. x 1/2.

Figs. 6, 9-12. Colonies with massive base and lamellate processes. Fig. 6: Lagoon
Pond, Martha’s Vineyard. YPM +940. x 1/2. (Viewed from side.)
Fig. 9: Double Beach (Branford); Conn. YPM) 222019 ei
(Viewed from side.) Figs. 10, 11, 12: Bradley Point, West Haven,
Conn. YPM 72026B. x 1/2. (Viewed from above.)

Fig.

Fig.

Fig.

Fig.

PLATE 4
Variations of colony form in Halichondria bowerbanki
(All photographs from specimens preserved in alcohol.)

g. 1. Colony consisting of a mass of anastomosing, rounded branches; viewed

from side. Off Lighthouse Pt., New Haven, Conn. YPM #814. x 1/2.

. 2. Colony with chimney-like process; viewed from above. Off Lighthouse Pt.,

New Haven, Conn. YPM #814. x 1/2.

. 3. Encrusting colony with vertical processes; viewed from above. Double

Beach (Branford), Conn. YPM +1893D. x 7/10.

gs. 4, 5. Thin, encrusting colonies without processes; viewed from above.

Double Beach (Branford), Conn. YPM #1893D. x 7/10.

6, 7, 8. Colonies with thin, anastomosing branches; viewed from above.
Fig. 6: Double Beach (Branford), Conn. YPM 3840. x > 1/2. Figs.
7, 8: Double Beach (Branford), Conn. YPM #1911E. x > 1/2.

Variations of colony form in Microciona prolifera.
(All photographs from dried specimens.)

g. 9. Large colony consisting of an anastomosis of branches; viewed from side.

Off Lighthouse Pt., New Haven, Conn. YPM #815. x 2/5.

10. Colony with palmate branching habit; viewed from side. Off Mansfield
Pt. (East Haven), Conn. YPM 3808. x 1/2.

. 11. Encrusting colony with short vertical branches; viewed from above. Off

Mansfield Pt. (East Haven), Conn. YPM #808. x 1/2.

Lissodendoryx and Microciona
12. Lissodendoryx isodictyalis. Double Beach (Branford), Conn. YPM
+2096. x <3/5. (Alcohol)

13. Microciona prolifera. Lamellate colony. New Haven, Conn. YPM £262.
x 1/2. (Dried specimen)

14. Microciona prolifera. Cup-shaped colony. New Haven, Conn. YPM
+262. x 1/2. (Dried specimen)

Fig.

Fig.

Fig.
Fig.

Fig.

PLATES

. Haliclona canaliculata. Colony viewed from above. Double Beach (Bran-

ford), Conn. YPM #2017. x 2/3. (Alcohol)

. Haliclona canaliculata. Upper surface of a colony showing the characteris-

tic pattern of subdermal channels and openings from these into the
interior of the sponge. Dermal pores not visible. Double Beach
(Branford), Conn. YPM 31955B. x 6. (Alcohol)

. Isodictya deichmannae. Nantucket. YPM #2112. x 1/3. (Dried specimen)

. Isodictya deichmannae. Grand Banks. YPM #419. x 3/10. (Dried speci-

men)

. Isodictya deichmannae. Off Nantucket. YPM #2113. x < 1/2. (Dried

specimen)

Fig.
Fig.

Fig.
Fig.

Fig.

PEACE? 6

. Isodictya palmata. Kent Island, N.B. YPM #923. x 2/5. (Dried specimen)
. Isodictya palmata. Minas Basin, N.S. YPM #2114. x > 2/5. (Dried speci-

men)

. Isodictya palmata. Eastport, Me. YPM #151. x > 2/5. (Dried specimen)
. Isodictya deichmannae. Kent Island, N.B. YPM #890. x > 2/5. Wied

specimen)

. Isodictya deichmannae. Off New London, Conn. YPM #762. x > 2/5.

(Alcohol)

PLATE 7
Fig. 1. Haliclona oculata. Off Sandy Point, Block Isl., R. l. YPM #783. x > L/S:
(Dried specimen)

Fig. 2. Haliclona oculata. Off Sandy Point, Block Isl., R. I. YPM #782. x >1/3.
(Dried specimen)

PeARE8
Fig. 1. Haliclona oculata. Off Sandy Pt., Block Isl., R. I. YPM S791, x0 /3., (Dried
specimen)

Fig. 2. Haliclona oculata. New Haven, Conn. YPM #430. x 2/5. (Dried specimen)

Fig. 3. Haliclona oculata. Off Sandy Pt., Block Isl., R. I. YPM #784. x 1/3. (Drie
specimen) yet

PRATESS
Fig. 1. Haliclona oculata. Off Nantucket. YPM #2115. x < 1/3. (Dried speci-
mens)

Fig. 1A. Haliclona oculata. Off Woods Hole, Mass. 8-20 meters. YPM +126.
x <1/3. (Dried specimen)

Fig. 2. Haliclona oculata. Stellwagen Bank, Massachusetts Bay. 35-40 meters.
YPM #965. x 1/3. (Dried specimen)

Fig. 3. Haliclona oculata. Off St. Valéry-en-Caux, France. YPM #1073. x 1/3.
(Dried specimen)

Fig. 4. Haliclona oculata. Woods Hole, Mass. YPM #942. x 1/3. (Dried speci-
men)

Fig. 5. Haliclona oculata. Off Hammonasset, Conn. YPM 3855. x 1/3. (Dried
specimen)

PEALE 10
Fig. 1. Haliclona oculata. Portland, Me. YPM #2116. x <1/4. (Dried specimen)

Fig. 2. Haliclona oculata. Fipennies Ledge, Gulf of Maine; 75 meters. YPM
#995. x < 1/3. (Dried specimen)

Fig. 3. Haliclona oculata. Stellwagen Bank, Massachusetts Bay; 35-40 meters.
YPM #965. x 1/3. (Dried specimen)

PEAIGE
Variations of colony form in Haliclona loosanofft.
(All photographs from specimens preserved in alcohol.)

Figs. 1, 3, 8-11. Colonies with many oscular tubules arising from an encrusting
base; all viewed from above. Figs. 1, 3, 11: Milford Harbor, Conn.
YPM +859. x 3/4. Figs. 8-10: Bradley Pt., West Haven} Geum
YEM 2520251 x 2)/3:

Fig. 2. Colony with dichotomizing vertical branches; viewed from side. Milford
Harbor, Conn. YPM#859. x 3/4.

Figs. 4, 6. Colonies with long, horizontal branches partly fused to basal mass;
viewed from above. Hammonasset, Conn. YPM #1821. x 3/4.

Figs. 5, 7. Encrusting colonies with low oscular tubules; viewed from above.
Hammonasset, Conn. YPM #2023F. x 3/4.

Figs. 12-16. Colonies with long, thin, anastomosing branches; all viewed from
above. Fig. 12: Double Beach (Branford), Conn. YPM #1848. x 3/4.

Figs. 13, 14, 15: Double Beach (Branford), Conn. YPM #1838. x 3/4.
Fig. 16: Double Beach (Branford), Conn. YPM #1956D. x 2/3.

PEATE: '2

Fig. 1. Haliclona loosanoffi. Gemmules attached to the shell of a barnacle. Mil-
ford Harbor, Conn. YPM #614. x 10. (Alcohol)

Fig. 2. Haliclona loosanoffi. Colony with tall, vertical branches; viewed from side.
Solomons Isl., Md. YPM #627. x 4/5. (Alcohol)

Fig. 3. Haliclona loosanoffi. ‘Vypical colony from the Maryland population.
Solomons Isl., Md. YPM #627. x 4/5. (Alcohol)

Fig. 4. Haliclona loosanoffi. Gemmules from Solomons Island, Md. YPM +627.
x 10. (Alcohol)

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