The Burrowing Sponges

of Bermuda

t

KLAUS RUTZLER

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 165
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SMITHSONIA N CONTRIBUTION S TO ZOOLOGY • NUMBER 165

The Burrowing Sponges

of Bermuda

Klaus Rutzler

FEB151974

SMITHSONIAN INSTITUTION PRESS

City of Washington

1974
ABSTRACT

Riitzler, Klaus. The Burrowing Sponges of Bermuda. Smithsonian Contributions to Zoology, number 165, 32 pages, 26 figures, 1 table, 1974 The systematic*

and distribution of eight species of burrowing sponges found on the Bermuda platform are discussed. These sponges excavate limestone substrata to form burrows

in which they live, thus participating in processes of erosion and sedimentation.

Seven species belong to Cliona (Clionidae): C. caribbaea Carter, C. flavifodina,

new species, C. paucispina, new species, C. vcrmifera Hancock, C. dioryssa (Laubenfels), new combination, C. lampa Laubenfels, and C. amplicavata, new

species; one species belongs to Spheaosponfia (Spirastrellidae): S. othella Laubenfels. In addition to the three new species, two sponges are new records for

Bermuda: C. caribbaea (not sensu Verrill, 1907, and Laubenfels, 1950) and

C. vermifera. Among the C. lampa population three phenotypic forms are distinguished: formae lampa, occulta, and flavida. Appearance of live and preserved

specimens, internal structure, spicules, and excavations are described in detail

to permit field identification by nonspecialists. Scanning electron photomicrographs are used to clarify the fine structure of microscleres.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 4970. SERIES COVER DESIGN:

The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Riitzler, Klaus, 1936-

The burrowing sponges of Bermuda.

(Smithsonian contributions to zoology, no. 165)

1. Cliona. 2. Spheciospongia othella. 3. Sponges—Bermuda Islands. I. Title. II. Series:

Smithsonian Institution. Smithsonian contributions to zoology, no. 165.

QLI.S54 no. 165 [QL373.D4] 591'.08s [593'.46] 73-12864

For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402

Price 80 cents (paper cover)
Contents

Page

Introduction 1

Acknowledgments 2

Methods 2

List of Stations 4

Terminology 5

Family Clionidae Gray 5

Cliona caribbaea Carter 5

C. flavifodina, new species 9

C. paucispina, new species 12

C. vermifera Hancock 15

C. dioryssa (Laubenfels), new combination 17

C. I am pa Laubenfels 21

C. amplicavata, new species 26

Family Spirastrellidae Ridley and Dendy 29

Sphcciospongia othella Laubenfels 29

Literature Cited

 S 2

in

The Burrowing Sponges

of Bermuda

Klaus Riitzler

Introduction

Observant divers are generally impressed by the

abundance, diversity, and biomass of sponges in

many shallow-water habitats throughout the tropical western Atlantic. Burrowing sponges are the

least conspicuous, and their importance in processes of erosion, sediment production, and calcium

carbonate dissolution has been greatly underestimated in the past (Goreau and Hartman, 1963;

Neumann, 1966; Riitzler, in preparation). Most

limestone excavating sponges belong to the family

Clionidae, but members of Spirastrellidae and Adociidae are also known to burrow (Riitzler, 1971;

Pang, in press; Riitzler and Rieger, 1973). In fact,

the concept of the Clionidae as a family, defined

by their burrowing habit, requires redefinition as

more material and data become available. Generic

and familial placement of species should be reconsidered from the standpoint of recognized procedures of sponge systematics, rather than separating

burrowing and nonburrowing forms.

The purpose of this paper, however, is not systematic revision, particularly not of higher taxa.

Having visited Bermuda during several consecutive years, it became clear to me that manuals describing the local flora and fauna are urgently

needed. During its seventy years of existence the

Bermuda Biological Station for Research has been

a center for investigations in every discipline of

nature sciences, including the systematics of local

plants and animals. Presently, there is an increasKlaus Riitzler, Department of Invertebrate Zoology, National

Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

ing emphasis on educational activities at the station, most of which, as can be expected at this

location, center on marine organisms. Whether

studies and research aim at embryology, physiology,

and ecology or at calcification, bioerosion, and

sedimentation, the organisms are still the key material, and knowledge of their kinds, occurrence,

and distribution is of first importance. Some taxa,

of course, are better known than others, but most

local systematic treatments need to be corrected

and updated with new findings. Also, systematic

papers, like most technical communications, tend

to be written for a small number of specialists and

are not readily understood by the less specialized

student. Many morphological and anatomical features that are essential for classification are difficult

to describe in words, and printing costs have generally curtailed illustrations.

This contribution is an attempt to overcome

some of the aforementioned problems and to describe and illustrate a small but significant fauna.

I hope that a person untrained in sponge systematics can, by using this report, either identify

a given excavating sponge from Bermuda, or determine with reasonable certainty that it had been

overlooked during my surveys.

Until recently, no study has ever been devoted

entirely to the systematics of burrowing sponges in

any one area of the tropical western Atlantic. Pang

(in press), describing excavating sponges from

Jamaica, increased the number of species known

from that island from two to thirteen. Since her

paper was still in press during preparation of this

manuscript, no discussion of her results is possible

in the systematic section.
Among the few publications on Bermuda

sponges, only two include descriptions of burrowing sponges. Verrill (1907) described Heterocliona

cribaria (Schmidt). Although this species has to

be considered a synonym of Spheciospongia othella

Laubenfels, Verrill was correct in suspecting it of

having a burrowing stage. Verrill (1907) also described the massive stage of what he took to be

Cliona caribbaea Carter. He failed to find microscleres and, with second thoughts about his identification, proposed the name Cliona sordida in a

footnote. Ever since, reports of massive "Cliona

caribbaea" have appeared in the literature (e.g.,

Laubenfels, 19S6, 1950, 1953; Wells et al., 1960).

1 have not found this massive sponge during my

work, and there is no evidence that Cliona caribbaea attains a massive (gamma) stage, like Cliona

celata Grant or Cliona viridis (Schmidt) (Topsent, 1900), and it is certainly not conspecific with

Cliona viridis. It is quite possible that Cliona sordida, if a Cliona at all, is a valid species. Laubenfels (1950) described massive cake-shaped "Cliona

caribbaea," abundant at Daniel's Head, off the

west end of Bermuda. He found it nowhere else

and speculated that his Cliona lampa had taken

over its ecological niche since VerriU's paper. This

might be a little far fetched but the fact is that

during the present study, even after two extensive

diving surveys, none of the bright yellow-ocher

massive sponges have been found. It is curious,

also, that a naturalist like Verrill should have

overlooked an abundant and conspicuous sponge

like Cliona lampa. My own related observations,

and recent findings in Florida (A. Antonius, personal communication), indicate that the conspicuous incrusting growth form of this species could

be the result of drastic environmental events that

occurred after Verrill's time. More work is planned

on this subject. In addition to the two species just

mentioned, and to Spheciospongia othella, Laubenfels (1950) described Spirastrella dioryssa. He

suggested the possibility that this species could be

a Cliona. Although the excavating habit of this

sponge could be confirmed during the present

study, the species is transferred to Cliona on the

basis of the type and structure of its skeleton.

This report adds five burrowing sponges to the

list, three new species and two new records for

Bermuda, one of which is considered the "true"

Cliona caribbaea Carter. The illustrations are inSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tended to be comprehensive and representative.

Scanning electron photomicrographs aim to clarify

details that cannot be resolved by light microscopy,

but they are not a substitute for procedures that

can be repeated with standard optical laboratory

equipment. I advise using the figures like a taxonomic key to narrow down the possibilities of

identification, but not to rely on any one or two

agreeing features because variability and overlap

of characters can mislead even the most experienced. Text and measurements should always be

consulted before a final decision is reached.

ACKNOWLEDGMENTS.—Financial support for the

project was provided by Smithsonian Research

Foundation Fund 427242. I thank Mr. W. R.

Brown and Miss M. J. Mann for operating the

scanning electron microscope. This is Contribution No. 575, Bermuda Biological Station.

METHODS.—Fieldwork for this study was conducted between October 1969 and January 197S.

All specimens were collected by the author, by

wading, breath-held or scuba diving, using hammer arid chisel. In the laboratory, sponge-infested

substrata were broken down and studied under

low-power microscope. Small, dean, separate

samples of epilithic and endolithic (ecto- and

choanosomal) sponge tissue were taken for spicule

mounts. Care was taken to avoid contamination

by other sponges invading the same substratum

fragments.

Quick preparations which assure minimum loss

of minute spicules are best made by dissolving

tissue fragments, placed on a microscope slide in

a drop of commercial bleach (5-10 percent sodium

hypochlorite). Small tissue pieces, dehydrated and

mounted in a medium like Canada balsam, are

also useful for locating small elements of the skeleton. For permanent slides, clean spicules are obtained by boiling tissue in concentrated nitric

acid, rinsing in water, and dehydrating in alcohol.

For scanning electron microscope viewing, clean

spicules were dried on a small circular cover slip

and coated with 20 nm of gold. A Cambridge

Stereoscan Mark II A was used at X 200 - X 5000,

rarely at X 10,000 or X 20,000 primary magnification.

To reveal the structure of burrows, substratum

pieces are best cleaned of organic material by soaking overnight in sodium hypochlorite solution. For

study of anatomy and histology, two methods have
NUMBER MS

been used successfully. For both, the material was

fixed in buffered 4 percent formalin-seawater (pH

7-7.5), which was replaced by 80 percent ethyl

alcohol after not more than 2 weeks. Method 1 involved decalcification in several changes of 5 percent nitric acid, neutralization in 5 percent sodium

sulfate solution for 12 hours, and rinsing in running up water for 6 hours. The tissue was then

embedded either in paraffin for standard microtome

sectioning, or iu 12 percent gelatine for sectioning

frozen by a cryostate microtome. Toluidin blue,

Mallory's tripple stain, or Ehrlich's hematoxylineosin all gave good staining results. For Method 2,

fragments of substratum containing the sponge (up

to 2 cm*) were dehydrated in a graded series of

alcohol, soaked in three changes of propylene

oxide and vacuum embedded in Spurr Low Viscosity Embedding Media (Polysciences, Inc., Warrington, Pennsylvania, U.S.A.). Specimen blocks

were glued to microscope slides and wet-ground

and polished to 20 urn. The sections were stained

in 1 percent aqueous safranin (10-15 minutes) and

subsequently in 1 percent aqueous crystal violet

(5-10 minutes).

Measurements given in this report are ranges

(means in parentheses) for all specimens studied.

Means are omitted where less than 5 measurements

were made. Spicule sizes are based on 25 measurements per dimension for three representative

sponge specimens. The largest diameter was recorded for approximately circular structures, the

longest perpendicular axes for oval or irregular

features.

In the synonymy, species names and references

are restricted to those of the first describer and to

subsequent descriptions which, by their quality

and relevance, contribute most to the understanding of the species.

The material on which this report is based is

deposited in the National Museum of Natural Hiskm

FIGURE 1. Map of Bermuda showing collecting localities.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tory, Smithsonian Institution. Catalog numbers are

those of the old United States National Museum

(USNM). Additional reference specimens are deposited at the Bermuda Biological Station.

LIST OF STATIONS (Figure 1, Table 1).—The following is a list of collecting localities that were

visited during all surveys (Figure 1). Collecting

date and substratum information are given in the

material section of each species. Place names are

used as shown on the Bermuda Islands Road Map

1966 (Bermuda Department of Tourism and Trade

Development) and on chart H.O.27, U.S. Navy

Oceanographic Office. Burrowing sponge species

found at each station are listed in Table 1.

1. John Smith's Bay. Small reefs, 20 m off shore. 2 m.

2. Tobacco Bay. Rocks and small reef patches. 1-2 m.

3. Devonshire Bay. West flank, rocks halfway to point. Intertidal.

4. Coot Pond. Rocks at entrance. Intertidal to I m.

5. Castle Harbour North. Patch reef between Blue Hole and

Navy Base Peninsula. 15-3 m.

6. The Narrows. Ship channel, east off Town Cut. Patch

reef. 4 m.

7. Church Bay, Harrington Sound. Subtidal parts of caves

and rocks. Intertidal to 1 m.

8. Ferry Reach, next to Whalebone Bay. Loose rocks; intertidal.

9. Whalebone Bay. East flank and rocks at entrance. Intertidal. tide pools, to 1 m.

10. Ferry Reach. Rocks at "Barnacle" Bay entrance near

Biological Station. Intertidal to 15 m.

11. Three Hill Shoals. Patch reefs. 3-10 m.

12. Castle Harbour East. Patch reef between Tucker's Town

and Navy Base Peninsula. 05-25 m.

13. Church Bay. South Shore. Boiler Reefs. 05-4 m.

14. Harrington Sound. Rock west off Trunk Island. 05-15 m.

15. Castle Harbor West. Off Waliingham Bay. 05-1 m.

16. Harrington Sound. Northeast comer of Hills Island.

Small caves. 05-15 m.

TABLE 1.—Distribution of burrowing sponge species found at collecting stations

in Bermuda

Stations "8

1. John Smith's Bay

2. Tobacco Bay

3. Devonshire Bay

4. Coot Pond

5. Castle Harbour North

6. The Narrows

7. Church Bay, Harrington Sound

8. Ferry Reach

9. Whalebone Bay

10. Ferry Reach

11. Three Hill Shoals

12. Castle Harbour East

13. Church Bay, South Shore

14. Harrington Sound

15. Castle Harbour West

16. Harrington Sound ,

17. North Rock

18. Daniel's Head

19. Battery Bay

20. North Rock South

21. Ferry Reach Entrance

22. Harrington Sound

x

X X

X X

X

X

X

X

X X

X X

X X

X X

X

X X

X

X

X X

X

X X

X X

X

X

X X

X X

X X

X

X X

X X

X

X X

X

X

X

X

X

X

X X

X

X X

X X

X

X X
NUMBER MS

17. North Rock. Reef*. 3-7 m.

18. Daniel'* Head. Patch reeft, 0.5-3 m.

19. Battery Bay. Boiler Reefs. 6-12 m.

20. North Rock South. Patch reef 300 m south of light tower.

5-8 m.

21. Ferry Reach Entrance, North Shore. On and between

railway pillars. 3-5 m.

22. Harrington Sound. Cockroach Island, west of Abbot's

Cliff. Rocks, 1 m.

TERMINOLOGY.—A minimum of technical terms

has been used, most of which can be found in

Borojevic* et al. (1968). Designations for spicule

types become dear with a glance at the illustrations. Terms like "head." "neck," and "shaft" for

sections of the pin-shaped tylostyles are selfexplanatory. The general structure of a dionid

sponge has been described and illustrated by Goreau

and Hartman (1963). Ectosome is the peripheral

region of a sponge lacking choanocyte-chambers;

choanosome contains choanocyte-chambers. In most

burrowing sponges, the major part of the sponge

body is inside the rock and only parts of the

ectosome show on the substratum surface. These

are called papillae and bear ostia or an osculum (openings of the incurrent and excurrent

canals). The perforations on the substratum

surface through which they protrude are called

papillary perforations, which can be extended by

a papillary canal or lead directly into the main

burrow. The burrow can have a honey-combed

appearance. The units are then called chambers

(not to be confused with the choanocyte-chambers

of the tissue), which are separated by walls but

are in communication through small holes, foramina.

Family CLIONIDAE Gray

CUona caribbaea Carter

Fionas 2-4

CUona caribbaea Carter, 1882:346-347, fig. 26_Topsent,

1889:49.

CUona viridu_Topsent, 1900:84-98 (in part), pi. 3: fig. 3d.

CUona viridis var. caribbaea.—Topsent, 1932:563-565, fig. 5.

DESCRIPTION.—Ectosome: The papillae (Figure

2a) exhibit basically a greenish color. Specimens

can be grayish ocher or dark brown with a green

tinge; others arc light olive or yellow olive; the

majority is of dark olive color. In alcohol, the

chlorophyll of the symbiotic algae dissolves and

leaves the papillae gray. Dry specimens are grayish

ocher. The papillae are circular to oval, and are

usually spaced 1-5 mm from each other, but they

have a strong tendency to fuse. Their diameters

range from 0.5 X 0.5 mm to 4.0 X 3.7 mm (2.4

X 1.9 mm). Fused complex papillae are 3.0 X

1.5 mm-22.0 X 9.0 mm (7.2 X 3.0 mm). Very

seldom fusion goes as far as to form continuous

incrustations, to the extent of 2-3 cm2. The following observations can only be made on undisturbed specimens underwater. The edges of the

papillae are raised to about 1 mm above the substratum; their center is depressed. Some of the

larger ones bear one or, at most, two oscula with

a raised rim (1-2 mm), which is always lighter in

color (yellowish) than the papilla. Diameter of

the osculum depends on the activity of the animal;

it can reach 1.2 mm. Ostia are present on all

papillae and can be opened to 0.3 mm. Intracellular symbiotic algae (zooxanthellae) (Figure 2/)

are packed in rows between the perpendicular

spicule strands. They are spherical and measure

8-10 urn in diameter. They decrease in number

with distance from the papillary surface. There is

also a well-developed connective tissue with numerous fusiform contractile cells, which account

for the strong contractibility of the papillae.

Choanosome: Breaking up the substratum exposes sponge tissue of rich yellow ocher color that

turns to pale gray in alcohol, but changes little

upon drying. The tissue fills the ill-defined, irregularly elongate chambers (Figure 26) except for the

major excurrent canals, measuring 0.5-1.0 mm in

diameter. The chambers are interconnected by

tissue strands of 0.1-1.5 mm. Interstitial spaces in

very porous substrata are also infiltrated. Irregular

or spindle-shaped cells, 24 X 7 um-30 X 10 urn,

occur in small accumulations. They contain 1 um

basophilic granules (gray cells). Some specimens

collected in January contained egg cells and sperm

cysts in various stages of development. Choanocytechambers measure 30-35 um.

Excavations: Single papillary perforations (Figure 2c) measure 0.5-3.2 mm (1.5 mm). Chambers

start at 0.5-2.0 mm under the substratum surface.

The chambers (Figure 2d) are irregular, ragged,

and frequently fused, particularly in porous substrata. They measure approximately 1.9-7.0 mm

X 1.3-2.9 mm (4.0 mm X 2.0 mm) if they are well
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—Cliona caribbaea: a, papillae of live specimen; b, tissue penetrating substratum; S

papillary canals in left upper coiner; c, papillary perforations; d, chambers inside pclccypod

shell; e, vertical section through papilla showing excurrent canal, arrangement of tylostyles and

distribution of zooxanthellae; /, intraccllular zooxanthellae enlarged. (Magnification: a-d, X 3;

e, X 50; /, X 850.)
NUMBER 165

FIGURE S—Cliona caribbaea, spicules: a, lylostyles; b, tylostyle head; c, d, spirastcrs and tylostyle

beads. (Magnification: a, X 140; b-d, X 850.)

defined. There are 6 to at least 15 foramina per

chamber, which are approximately circular and

0.2-1.2 mm (0.5 mm) in diameter. The thickness

of the separating walls varies between 0.2 and 1.0

mm. Excavations reach 15 mm into the substratum.

Spicules: Tylostyles occur everywhere, but show

orientation perpendicular to the substratum only

in the papillae (Figure 2e). Spirasters are rare or

absent in the papillae, moderately abundant in

the choanosome. Tylostyles (Figures Sa-d, 4a) are

straight or slight bent. The point is sharp and

gradually tapering from the middle of the spicule.

The heads are well set off and usually spheroid.

If they are ovoid, the longer axis of the head tends

to be perpendicular to the axis of the shaft. Deformations of the heads are common, but rarely

are there annular swellings behind them on the

shaft. The spirasters (Figures 3c, d, 4b, c) are very

delicate and easily overlooked. Under the light

microscope they appear as finely spined wavy lines,

the majority with 4-7 bends. Some are straight or

W-shaped.

Spicule dimensions (in um): Tylostyles, length

X width: 204.1-410.0 X 2.6-8.1 (319.0 X 6.0).

Neck width: 1.5-7.4 (4.6). Head length X width:

7.4-13.7 X 7.0-14.8 (9.7 X 10.3). Spirasters,

length X width (including spines): 9.6-38.4 X

0.8-3.0 (32.8 X 1.8). Spine width: not resolvable

by light. Number of spines: 18-50 + (40 +).

Number of bends. 3-10 (5.9).

REMARKS.—This species has been greatly misinterpreted in the sponge literature of the tropical

western Atlantic. Carter (1882) and Topsent (1900,

1932) accurately illustrated the characteristic spirasters. There are indications in the Bermuda

material that Cliona caribbaea might adopt an incrusting habit, at least to a certain extent. Bermuda

specimens have smaller tylostyles than the holotype

from St. Vincent, West Indies. Also the depressed

form of the tylostyle heads is more obvious in the

Bermuda material. Three specimens studied for

comparison from Dominica (West Indies, close

to the type-locality) and Galeta Island (Panama)

have tylostyles exactly as figured by Carter (1882),

with mean tylostyle dimensions, length X width:

399.0 X 9.6 jim (Carter: 396 X 10 um). Massive

(gamma stage) specimens without spirasters, as

described by Verrill (1907) and Laubenfels (1950)

for Bermuda, and specimens (of various authors)

with spirasters bearing prominent spines are almost

certainly not conspecific. The latter includes material described by Hechtel (1965) as Cliona viridis

(Schmidt), and placed in synonymy with Cliona

caribbaea.

MATERIAL.—USNM 24346; Station 5; 31 December 1969; in dead coral and pelecypod shell. USNM

24347; Station 11; 13 January 1970; in dead parts

of coral (Millepora, Porites). USNM 24348; Station

21; 15 June 1972, in pelecypod shell. Station 2; 8

May 1971; in dead parts of coral (Dichocoenia).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICURE 4.—Cliona caribbaea, spiculcs (SEM): a, tylostylc heads; b, c, spirasters and derivatives.

(Magnification: a, b, X 1800; c, X 2700.)
NUMBER ltt

Station 5; 31 December 1969; in dead coral. Station

5; 12 January 1970; in dead coral. Station 5; 30

December 1971; in rock. Station 9; 30 April 1971;

in coralline incrusted rock. Station 10; 31 May

1972; in rock. Station 12; 27 April 1971; in pelecypod shell, rock, dead parts of coral (Oculina). Station 17; 16 May 1971; in dead coral. Station 17;

9 June 1972; in rock. Station 18; 13 May 1971; in

dead coral (Agaricia). Station 19; 7 June 1972; in

dead coral. Station 21; 12 January 1973; in dead

pelecypod shell and coral (Oculina).

DISTRIBUTION.—With certainty only tropical western Atlantic.

Cliona flavifodina, new species

FIGURES 5-7

DIAGNOSIS.—Papillae yellow-brown, circular, discrete, 2.8 mm (mean diameter). Excavations irregular, tunnel shaped, confluent; filled by yellow,

tough tissue. Tylostyles straight, with dropletshaped or elongate heads, 318.1 X 7.4 (Am (mean

length X width). Spirasters with thin shaft, average 3-4 bends, 36.9 X 1.6 um (mean length X

width of shaft), and discrete slender spines, 4.5 urn

(mean length).

DESCRIPTION.—Ectosome: The color of the pap

illae varies from bright yellow to grayish yellow

and chestnut brown. In alcohol and upon drying,

it fades to pale gray or grayish brown. The papillae are inconspicuous, level with the surface of the

substratum, and well spaced (Figure 5a). Not

more than ten could ever be observed in any one

specimen. Their shape is circular, rarely elongate;

their diameters are 1.5 X 1.5 mm-5 X 3 mm (2.8

X 2.2 mm). Papillary fusion is an exception. The

surface of the papillae is irregularly porous. Oscula

are rare; they open to about 3 mm diameter and

have a slightly raised rim. The ectosome contains

an abundance of granular cells, ovoid to irregularly

elongate, 16 X 16 um-38 X 8 um, with granules

of 1 um. A small amount of zooxanthellae was

observed in one specimen.

Choanosome: Grayish yellow tissue lines the

papillary canals. These open into large, irregular

spaces filled with a tough cavernous tissue (Figure

5b). The color of the choanosome is bright yellow

to deep ocher, occasionally with an orange tinge.

In alcohol or when dried, it becomes pale gray.

In addition to the granular cells mentioned above,

there are numerous cells of a second kind. They

are more or less spherical, 18-22 um in diameter,

with a small anucleolate nucleus (like the gray

cells), and packed with large spheroid basophilic

granules of 1.5-2 um (Figure 5/).

Excavation: The size of the circular papillary

perforations (Figure 5c) corresponds to that of the

papillae, although in the macerated specimens

measured, it only ranged from 0.5 X 0.5 mm-3.2

X 2.2 mm (1.7 X 1.1 mm); one double perforation

was 4.8 X 4.3 mm. The papillary canals are 1.1-7.0

mm (3.1 mm) long, until they widen and reach

the choanosomal portion of the sponge. No chambers are developed, at least in the present material.

There are large irregular spaces (Figure 5d) with

tapering tunnels radiating into the substratum to a

depth of 20 mm. Horizontal extension of the excavations reaches 25 mm.

Spicules: The papillae are armed with perpendicular typostyles (Figure be). The choanosome

contains tylostyles and large numbers of spirasters.

The tylostyles (Figures 6a-d) are mostly straight;

occasionally they are bent shortly behind the head.

The heads are droplet shaped or elongate ovoid;

most of them show a central dark spot in transmitted light. Secondary swellings produce characteristic violin-shaped and almost rectangular heads,

with an opaque spot close to the neck. The spirasters (Figures 6b-d, 7) are uniform in appearance,

with a thin shaft and a moderate number of slender

but prominent spines. Most of them have four

bends, but some are straight with reduced spines

(amphiasters).

Spicule dimensions (in um): Tylostyles, length

X width: 178.0-409.4 X 0.7-11.1 (318.1 X 7.4).

Neck width: 2.2-10.4 (5.8). Head length X width:

5.9-24.1 X 3.7-17.8 (15.5 X 11.5). Spirasters,

length X width (shaft only): 14.4-^5.6 X 0.5-3.2

(36.9 X 1.6). Spine length X width (at base):

1.4-7.7 X 0.6-1.9 (4.5 X 1.5). Number of spines:

8-27 (17.4). Number of bends: 1-5 (3.3). Amphiasters, length X width (shaft only): 9.6-38.4 X

0.6-2.5 (22.9 X 2.3). Spine length X width (at

base): 1.8-3.2 X 0.5-1.6 (2.7 X 1.1). Number of

spines: 5-14 (9.3).

REMARKS.—A number of Cliona specimens attributed in the literature to C. caribbaea or C.

iriridis could in fact belong to the species described

above. This view is particularly supported by
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 5.—Cliona flavifodina: a, 2 papillae (arrows) of live specimen; b, papillary canals and

tissue inside substratum; c, papillary perforations; d, papillary canal (arrow) and large burrow

(black area, to the right of embedded vermetid tube); e, vertical section through papilla; /,

choanosomal granular cells. (Magnification: a-d, X 3; e, X 50; /, X 520.)
NUMBER 165 11

U

a

FIGURE 6.—Cliona flavifodina, spicules: a, tylostyles; b-d, spirasters and tylostyle heads.

(Magnification: a, X 140; b-d, X 850.)

FIGURE 7 Cliona flavifodina, spicules (SEM): spirasters. (Magnification: X 1800.)
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Hechtel's (1965) account of two specimens of C.

viridis from Jamaica. The details of his description

of papillae, color, structure, and spicules agree well

with my observations on C. flavifodina, the only

exception being that Hechtel's specimens formed

small galleries (1-3 mm in diameter) instead of

extensive tunnels. Future studies will certainly

reveal the taxonomic significance of this difference.

This species, as judged from its spicules, is related to C. caribbaea. It differs by the shape of the

tylostyle heads, by size, spination, and number of

bends of the spirasters, by the shape of the burrow, and by the lack of zooxanthellae.

The species name (Latin, flavus: yellow; Latin,

fodina: pit, mine) refers to the appearance of the

tissue-filled burrow.

MATERIAL.—USNM 24353 (holotype); Station

11; 13 January 1970; in coralline incrusted dead

coral. USNM 24354 (paratype); Station 5; 12 January 1970; in dead coral. USNM 24352 (paratype);

Station 13; 28 April 1971; in dead coral. Station 5;

12 January 1970; in pelecypod shell. Station 5; 3

January 1972; in rock. Station 7; 6 January 1970;

in rock. Station 10; 31 May 1972; in rock. Station

17; 9 June 1972; in dead coral. Station 19; 7 June

1972; in dead coral. Station 21; 21 January 1973;

in dead coral.

DISTRIBUTION.—Bermuda.

Cliona paucispina, new species

FIGURES 8-10

DIAGNOSIS.—Brown incrustations, penetrating

rock crevices, limited excavating efficiency. Densely

arranged, slender, bent tylostyles, with inconspicuous, frequently subterminal heads, 340.7 X 6.2 urn

(mean length X width). Spirasters with thin shafts,

average 4-5 bends, 28.8 X 1.3 um (mean length X

width of shaft), and low spines, 1.6 um (mean

length). Spiraster spines on shaft characteristically

reduced. Abundance of intracellular zooxanthellae.

DESCRIPTION.—This sponge forms irregular,

patchy, thin incrustations (Figure 8a) on concave

surfaces of rock. Judging from the small extent of

penetration, it is not a very effective burrowcr.

The erosion pattern on the limestone surface suggests, however, that not only existing crevices are

penetrated but that active excavation occurs. The

sponge crusts are about 0.5-2.5 mm thick and S

X 3 mm—10 X 35 mm in horizontal extension.

Maximum area coverage observed was 7.5 cm2.

Ectosome: The color at the smooth surface of

the incrustations is chestnut brown to grayish

brown in life, with small yellowish areas around

the oscula. The color in alcohol is drab. The oscula

are slightly elevated and open to 6 mm in diameter.

No observations on open ostia have been made.

Zooxanthellae are present in the tissue in large

numbers. They are intracellular and measure 10

um, like in Cliona caribbaea. There are also some

of the large cells with inclusion described below.

Choanosome: It is light brown to yellowish

brown, turning to grayish brown in alcohol. The

tissue penetrates and fills small irregular spaces

in the substratum (Figure 8b). Larger spaces are

only lined with tissue. Its horizontal extension does

FIGURE 8.—Cliona paucispina: a, ectosomal incrustation with osculum partly contracted;

b, tissue filling burrow. (Magnification: a, b, X 3.)
NUMBER 165 13

not exceed the extension of the incrustations on

the surface. Depth of penetration reaches 12 mm

at the most. Zooxanthellae are still present here.

Large cells, elongate fusiform or irregular in outline, which measure 30 X 10 um - 40 X 24 um, are

very conspicuous. They are packed with weakly

staining spherical inclusions of 1.5-2.4 um. The

nucleus of 3 um is provided with a nucleolus.

Choanocyte-chambers measure 30 um in diameter.

Excavations: Little can be added to the observations above. The substratum surface below the

incrustations is eroded and existing spaces are

enlarged. There is no chamber formation, nor distinct tunnels. All observations were made on very

porous or jx>orly cemented substrata which can

easily be broken down. Future findings may demonstrate that this sponge can produce more distinctive burrowing patterns in compact substrata.

Spicules: A series of vertical tylostyles at the

surface is also present in this species, although this

structure can be obscured in contracted specimens.

The spicules are densely packed throughout the

tissue; a small number of spirasters is interspersed

among the tylostyles. Some tylostyles are organized

in rows which branch and brush out toward the

surface The tylostyles (Figures 9a,b,e, 10a) are

long and slender. They are usually bent in the

upper third or fourth of their length, behind the

head. The heads are not well set off, narrow, and

elongate. Most of them are mucronate or subterminal; many are reduced. The spirasters (Figures

9fl-e, 106) are delicate, with slender shafts and

small spines. The number of spines is frequently

reduced on the shaft, but small spines are always

w

FIGURE 9.—Cliona paucispina, spicules: a, tylostyles and spirasters; b, e, spirasters and tylostyle

heads; c, d, spirasters. (Magnification: a, X 140; b, e, X 520; c, d, X 850.)
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 10.—Cliona paucispina, spicules (SEM) : a, tylostyle head; b, spirasters and derivatives.

(Magnification: a, X 1800; b, X 2700.)

present at the ends of the spicule. Some shafts are

straight or irregularly bent, and W-shaped forms are

not uncommon.

Spicule dimensions (in urn): Tylostyles, length

X width: 124.6-471.7 X 3.0-8.9 (340.7 X 6.2).

Neck width: 2.2-7.4 (5.5). Head length X width:

5.2-22.2 X 4.1-12.6 (12.8 X 8.9). Spirasters, length

X width (shaft only): 12.8-43.2 X 0.8-3.0 (28.8

X 1.3) Spine length X width (at base): 0.5-3.2

X 0.5-1.9 (1.6 X 0.8). Number of spines: 6-25

(14.9). Number of bends: 0-7 (4.4)

REMARKS.—In spite of the similarity of spicule

complements between this and the two preceding

species, there is little likelihood of confusion.

Cliona paucispina is well characterized by the incrusting, predominately epilithic habit, by the

dense arrangement of tylostyles, and by the reduced spination of the spirasters.

The species name is derived from Latin paucus:

few, little, and spina: thorn.

MATERIAL.—USNM 24349 (holotype); Station 5;

12 January 1970; in dead coral. USNM 24350

(paratype); Station 13; 28 April 1971; in dead

coral. USNM 24351 (paratype); Station 5; 30 December 1971; in dead coral. Station 2; 28 October

1969; in rock.

DISTRIBUTION.—Bermuda.
NUMBER 165 15

Cliona vermifera Hancock

FIGURES ii-!3

cliona vermifera Hancock. 1867:259-240. pi. 8: fig. 2.—Toptent. 1889:35*—Hcchtei. 1965:60-61.

DESCRIPTION.—Ectosome: The papillae (Figures

lla, b) are usually of vivid orange-red color, light

ocher in alcohol, or dry. In a few specimens they

afe dark brownish orange. Their shape is circular

or oval, sometimes rather irregular. They are about

•%

FIGURE 11.—Cliona vermifera: a, b, expanded and contracted papillae of live specimen; c, papillary canals and burrows in coral rock; d, papillary perforations; e, chambers with large

foramina in pelecypod shell; /, vertical section through papilla; g, choanosomal a-granular cells.

(Magnification: a-e, X 3; /, X 140; g, X 850.)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FicutE 12.—Cliona vtrtnifera, spicules: ; tylostyles and microsderes; b, tylostykt;

c, d, microsderes (smooth spinsters). (Magnification: a, b, X 140; e, d, X 520.)

FIGURE IS.—Cliona vermifcra, spicules (SEM): Microsderes

(smooth spirasters). (Magnification: X 1500.)

2 mm apart, with little tendency to fuse and measure 0.3 X 0.3 mm-1.5 X 2.5 mm, 1-1.5 mm for the

majority. Oscula are under 1 mm in size, with a

raised yellow-orange rim. Ostia can expand to

0.3 mm; their average size, however, is under 50 urn.

Choanosome: The large chambers are almost

filled with fleshy, deep red, pale orange, or bright

orange tissue that fades to light brown in alcohol

or when dried. The same weakly staining, spheroid,

a-granular cell type described for Cliona amplicavata is present here in large numbers (Figure

llg). They are particularly accumulated near the

substratum walls of the chambers, in the diaphragm

filling the chamber foramina, and also in strands

leading through the choanosome. Choanocytechambers measure 28 urn. Large spaces devoid of

cellular material can be noted in the center of the

gallery chambers and to both sides of the foramina.

Excavations: The papillary perforations (Figure lid) are very small and can easily be confused

with those of Cliona lampa forma occulta: 0.3-1.8

mm (1.0 mm). However, there are well-developed

papillary canals, 1.3-3.5 mm (2.0 m) long, and

large chambers. The chambers (Figures lie,*) are

basically ovoid, the longer axis parallel to the substratum surface and with somewhat angular walls.

Their size (length X width X height) is 2.7 X

2.7 X 1.0 mm-77 X 4.8 X 2.0 mm (4.8 X 3.5 X

1.4 mm). They are separated by walls of 0.3-1.9

mm. The walls are perforated by at least six large

foramina, 0.3-1.1 mm (0.6 mm) in diameter. The

galleries extend 20 mm into the substratum.

Spicules: Tylostyles in the papillae are placed

perpendicular to the surface (Figure 11/). Elsewhere they are scattered or loosely organized into

strands. There appear to be two size classes, one

shorter and thicker than the other, but neither has

a particular location and there are many transitional forms. The tylostyles (Figures 12 a,b) are

mostly straight; some are bent in the third behind

the head. Frequently, they do not taper toward

the point until the last fourth of their length. The

heads are spheroid but mostly mucronate, occa-
NUMBER 165 17

sionally subterminal. The raicroscleres (Figures

12 a,c,d, 13) occur everywhere in the tissue but

they are more abundant in the choanosome. They

are smooth spiralled or undulated rods of uniform

thickness throughout their length. The ends are

rounded. These spicules are commonly called

"smooth spirasters."

Spicule dimensions (in um): Tylostyles, length

X width: 133.5-115.0 X 3.0-8.9 (242.4 X 5.7).

Neck width: 1.1-7.1 (4.6). Head length X width:

5.6-11.8 X 1.8-13.7 (10.7 X 8.9). Spirasters

(smooth), length X width: 16.0-76.8 X 1.0-4.8

(59.3 X 2.8). Number of bends: 0-9 (5.1).

MATERIAL.—USNM 24340; Station 9; 8 January

1970; in rock. USNM 24341; Station 13; 28 April

1971; in rock. USNM 24342; Station 18; 15 May

1971; in dead base of live coral (Mussa), dead

Millepora, pelecypod shell. Station 5; 31 December

1969; in gastropod shell. Station 5; 12 January

1970; in pelecypod shell. Station 7; 6 January 1970;

in rock. Station 10; 11 January 1970; in pelecypod

shell. Station 10; 31 May 1972; in coralline incrusted rock. Station 11; 13 January 1970; in

coralline incrusted coral rubble (Porites, Millepora). Station 12, 27 April 1971; in dead coralline

incrusted coral. Station 15; 3 May 1971; in rock.

Station 17; 9 June 1972. Station 18; 13 May 1971.

Station 19; 7 June 1972.

DISTRIBUTION.—Tropical and subtropical Atlantic and Indo-Pacific.

Cliona dioryssa (Laubenfels), new combination

FIGURES 14-16

Spirastrella dioryssa Laubenfels, 1950:98-99, fig. 44.

DESCRIPTION.—Ectosome: The ectosomal portions of the sponge which are showing on the substratum surface are of deep orange to yellow-orange

color. The color becomes light brown in alcohol

and after drying. Some of the smaller isolated structures (1.0 X 1.0 mm-6.0 X 3.5 mm) are approximately circular and resemble typical papillae,

beaiing oscula or pori. Commonly, however, there

are small irregular crusts with ragged outline that

have a tendency to fusion (Figure 14 a, b). The

crusts are about 1 mm high, 8X 4 mm-16 X 8 mm

in diameter. One large single incrustation measured 55 X 30 mm and covered an area of 5 cm2

.

Laubenfels (1950) reported specimens that were

"still not as much as 10 cm in diameter," with a

thickness reaching 3 mm. Oscula are irregularly

distributed over the incrustations. They too have

ragged edges; they measure 0.5 X 0.5 mm-2.0 X

3.0 mm and can sometimes be provided with a

minute membranous collar. Ostia are 0.1 mm in

diameter. There are densely accumulated granular

cells, irregular or spindle shaped, about 15 urn

long, with 1 um inclusions.

Choanosome: Distinct chambers, as well as crevices in porous substrata, are filled with pale orange

tissue that turns pale ocher upon preservation

(alcohol or dry). Granular (1 um granules),

spindle, or club-shaped cells (Figure 14g), probably the same kind noted above, occur in large

numbers. They are larger than those in the ectosome, 24 X 16 X 32 X 13 um, and it was observed that they show a remarkable resistance to

solution in sodium hypochlorite when spicule

mounts were prepared. Different stages of developing eggs measure up to 40 um (May); choanocytechambers measure 28 um.

Excavations: The size of the irregular depressions on the substratum surface (Figure 14d) corresponds with that of the tissue incrustations. In

macerated rock fragments containing the sponge,

they ranged 0.5 X 0.5 mm-8.0 X 5.6 mm (2.3 X 1.9

mm). The larger depressions are 4—6 mm deep and

lead directly into the chambers, very much like in

the incrusting form of Cliona lampa. Only small

papillae have a well-defined canal of 0.5-2.4 mm

(1.1 mm). The chambers (Figures \Acje) are

more or less spherical, but are not well defined

because the separating walls are frequently broken

down. They measure 1.1-3.2 mm (2.2 mm). The

walls are 0.1-1.0 mm (0.3 mm) thick and perforated by at least six foramina per chamber, 0.2-

1.2 mm (0.5 mm). Because of many perforations,

in addition to natural crevices, the walls are frequently reduced to single bars. Penetration into

the substratum attains 25 mm.

Spicules: A palisade of tylostyles is present in

the outer ectosome. Just below this, spicules occur

in a crisscross fashion (Figure 14/). In the choanosome they are sparse and without orientation,

some strands excepted. The tylostyles (Figures

15fl-c) are usually slightly bent. They have a slender,

gradually tapering shaft and a relatively large subspherical head, with a tendency to be mucronate.

Subterminal and reduced heads or a second swell-
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICURE 14.—Cliona dioryssa: a, incrusting portion of live specimen; b, S fusing papillae bearing

oscula; c, chamber arrangement in well-cemented limestone (pelecypod shell); d, papillary perforations and e, chambers in porous coral rock; /, vertical section through sponge crust; g,

choanosomal granular cells near tissue condensation (diaphragm) between 2 chambers. (Magnification: a-e, X 3; /, g, X 140.)
NUMBER 165 19

FIGURE 15.—Cliona dioryaa, spicules: a, tylostyles; b, tylostyle heads; c, spiraster and tylostyle

head; d-g, spirasters. (Magnification: a, X 140; b-g, X 850.)

ing behind the head are infrequent malformations.

The spirasters (Figures \bc-g, 16) are concentrated

in the ectosome without forming a cortex. They

are remarkably variable in size and shape and

many show a high degree of malformation. Although there are many transitional forms, two

types can be recognized. Type I is stout, with a

thick shaft densely set with large strong spines,

most of which are joined at the base. Type II is

more delicate, with a long slender shaft and small

spines, which are usually well spaced from each

other. Reduction of number or size of spines is

very common.

Spicule dimensions (in um): Tylostyles, length

X width: 106.8-391.6 X 3.7-7.4 (244.4 X 5.4).

Neck width: 3.3-7.0 (4.4). Head length X width:

3.7-17.8 X 1.9-11.5 (10.0 X 8.3). Spirasters I,

length X width (shaft only): 11.2-41.6 X 1.4-

4.8 (27.4 X 3.2) Spine length X width (at base):

1.6-6.7 X 1.0-4.0 (4.6 X 2.6). Number of spines:

7-26 (16.0). Number of bends: 0-5 (3.0). Spirasters II, length X width (shaft only): 19.2-43.2

X 0.6-2.2 (33.9 X 1.5). Spine length X width (at

base): 0.5-3.8 X 0.5-2.6 (2.3 X 1.3). Number of

spines: 7-24 (13.6). Number of bends: 2-6 (4.3).

REMARKS.—Laubenfels (1950) had some doubts

about the generic placement of his species. He

noted its penetration into the substratum and

stated that "if it is truly 'boring,' then it is a

Cliona, not a Sptrastrella at all." The burrowing

habit of Cliona dioryssa is confirmed here. However, it should also be noted that the relatively

small number of spirasters and the lack of an

ectosomal crust formed by spirasters are untypical

for the genus Spirastrella.

MATERIAL.—USNM 24343; Station 13; 28 April

1971; in dead coral. USNM 24344; Station 5; 3

January 1973; in dead coral and pelecypod shell.
FIGURE 16.—Cliona dioryssa, spicules (SEM) : Spirasters and derivatives. (Magnification: X 2700.)
NUMBER 165

USNM 24345; Station 10; 31 May 1972; in rock.

Station 5; 31 December 1969; in dead coral. Station 5; 12 January 1970; in rock. Station 5; 12

January 1970; in dead coral. Station 5; 3 January

1972; in rock. Station 5; 30 December 1971; in

rock. Station 6; 1 January 1970; in dead coral. Station 7; 6 January 1970; in dead coral. Station 9;

6 January 1970; in dead coral. Station 10; 11 January 1970; in rock. Station 10; 31 May 1972; in rock.

Station 11; IS January 1970; in rock, dead coral,

pelecypod shell. Station 13; 28 April 1971; in dead

coral. Station 14; 2 May 1971; in rock, pelecypod

shell. Station 16; 6 May 1971; in rock. Station 17;

6 May 1971; in coralline incrusted rock. Station 17;

9 June 1972. Station 19; 7 June 1972; in rock.

Station 20; 9 June 1972; in dead Millepora. Station 21; 12 January 1973. Station 22; 26 June 1972;

in rock.

DISTRIBUTION.—Bermuda.

CUona lampa Laubenfeb

Ficuus 17-20

CUona lampa LaubcnfcU. 1950:110-112. fig. 49.

DESCRIPTION.—Ectosome: This sponge has three

distinct morphological forms. Forma lampa (Figure 17a) is the one described by the original author.

It is also the most conspicuous variety. The ectosome encrusts the substratum with a continuous

very fine layer of vermilion, occasionally rusty red

tissue. It turns faintly gray in alcohol, light brown

in the dry state. The extension of the incrustations

ranges from 2 cm2 to approximately 1.5 m2 per

specimen. Oscula are scattered over the surface,

usually circular, 2-4 mm in diameter, with a raised

rim. Ostia measure about 0.8 mm. Forma occulta

(Figure 17b) is easily overlooked because only very

small circular papillae penetrate the rock surface

to the exterior. Their color is basically the same

as in forma lampa, perhaps less vivid. They are

densely spaced, about 25 per cm2, but fusion is

extremely rare. Diameter of the papillae is 0.2-

1.5 mm, mostly 0.8 mm. The larger ones bear circular oscula of 1-3 mm. The papillae are level

with the substratum surface; only expanded oscula

protrude. Forma flavida has the identical appearance as forma lampa, except that the incrustations

have sulfur yellow color. Only two yellow specimens have been found, one covering about 120

21

cm2, the other about 300 cm*. The histology and

cytology of this species have been discussed in detail

by Rutzler and Rieger (1973). An extensive contractile tissue, reaching 0.8 mm below the sponge

surface, is present in forma lampa.

Choanosome: The tissue completely fills small

galleries which are always developed if the sponge

burrows in compact substrata. In very porous substrata, as many of the invaded corals are, galleries

are ill-defined and existing spaces are permeated.

Color of the choanosome is dull rusty red, orange,

or yellow-orange in formae lampa and occulta,

grayish ocher in forma flavida. It fades to become

almost colorless after preservation. Specimens col*

lected in January and June contained numbers of

orange gemmules in the choanosome. They measure 0.7-0.9 mm, with a noncellular envelope 10 um

thick Between the enveloping membrane and the

reserve material filling most of the gemmule, there

is a thin layer of loosely spaced spindle-shaped

cells (Figure 186).

Excavations: The substratum surface that was

previously covered by the incrusting forms (lampa

and flavida) of this sponge is obviously eroded

(Figure 17c), but only to a certain degree, without

destroying major structural features. It looks

porous, like pumice stone, the pores leading directly into the chambers. At the location of the

oscula there are ragged depressions, 1.6-3.2 mm

(2.4 mm), about as deep as wide. They lead directly into the chambers. The sponge can infiltrate

porous substrata where chambers are obscure (Figure 17/). In compact substrata the small chambers

are spheroid, 0.5-2.0 mm (1.2 mm) (Figure 17c).

The walls of the chamber are 0.1-0.5 mm thick,

and pierced by numerous (at least four to nine)

foramina of 0.2-0.6 mm (0.3 mm). The substratum

can be penetrated to a depth of 80 mm. Form

occulta has small papillary perforations, 0.3-1.8

mm (0.8 mm) (Figure 17d). There is no distinct

papillary canal; the chambers start 0.1-0.6 mm

under the surface. The chambers are like in the

other forms, 1.0-2.0 mm (1.4 mm), perhaps somewhat more spaced, i.e., with thicker walls: 0.2-

0.8 mm. A maximum of 12 mm penetration into

the substratum was observed.

Spicules: In the ectosome, just below the surface, there is a layer of spiny microrhabds without

orientation and a series of perpendicular tylostyles

which partly penetrate the surface with their tips
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

.

 r 'id

•

... v

1

•

• •

FIGURE 17.—Cliona lampa: a, forma lampa incrusting the coral Diploria, 2 oscula are showing;

b, papillae of live forma occulta; c, eroded surface of pelecypod (Chama) shell burrowed by forma

lampa. Larger holes indicate former position of oscula; d, papillary perforations produced by

forma occulta; e, chamber arrangement in compact (pelecypod shell) and, /, in porous substratum

(coral). (Magnification: a-f, X 3.)
NUMBER 166 25

dS ^*

Ficunr 18 Cliona lampa: a, vertical section through incrusting tissue of forma latnpa;

b, section through gemmula. (Magnification: a, b, X 140.)

(Figure 18a). Below, there is a zone of tylostyles,

oxea, and mitrorhabds without orientation, but

rather parallel than perpendicular to the surface.

All three spicule types occur also in the choanosome but there they are quite rare. Some of the

tylostyles are organized into strands. The tylostyles

(Figure 19a) are straight and gradually tapering

to a sharp point. Their heads are spherical or

ovoid; the longer axis is always in the direction

of the shaft. The oxea (Figures 196, c, 20a, b, f)

are bent in the center and provided with minute

spines over the entire surface. At high magnification (scanning electron microscope), it can be seen

that the spines resemble hooks or double hooks.

Central swellings are very rare. The microrhabds

(Figures 19c, 20c-/) are mostly straight and densely covered by robust single or furcated spines.

Spicule dimensions (in um): Forma lampa.

Tylostyles, length X width: 118.4-259.0 X 0.8-

3.8 (196.7 X 2.8). Neck width: 0.8-3.2 (2.1). Head

length X width: 3.2-7.8 X 2.4-7.4 (5.6 X 4.3).

Oxea, length X width: 59.2-115.2 X 1.0-2.9 (85.7

X 1.8). Microrhabds, length X width: 4.0-25.6

X 0.8-3.8 (13.5 X 1.6).

Forma occulta. Tylostyles, length X width:

142.0-320.4 X 1.1-5.8 (245.6 X 3.7). Neck width:

1.0-3.7 (2.9). Head length X width: 4.0-8.0 X

1.9-8.0 (7.1 X 6.0). Oxea, length X width: 51.2-

94.4 X 0.8-5.6 (75.6 X 2.8). Microrhabds, length

X width: 6.4-21.6 X 1.3-4.8 (12.9 X 2.3).

Forma flavida. Tylostyles, length X width:

107.3-222.0 X 0.7-4.1 (183.8 X 2.8). Neck width:

0.5-3.7 (2.2). Head length X width: 3.7-7.4 X

3.0-6.7 (5.5 X 4.4). Oxea, length X width: 57.6

X 92.8 X 1.4-2.1 (73.2 X 1.7). Microrhabds,

length X width: 4.8-16.0 X 0.6-2.6 (12.4 X 1.8).

REMARKS.—Laubenfels (1950), in justifying the

naming of Cliona lampa, listed a number of characteristics to distinguish his new species from C.

vastifica Hancock. The present findings will weaken his argument considerably if it can be maintained that the three forms of C. lampa in Bermuda

are merely different phenotypes of the same

species: yellow and "rusty" red (rather than vermilion) color varieties are represented by formae

flavida and occulta (occasionally also lampa). Distinct burrows (chambers), circular in cross-section

(rather than permeation), are produced by all

forms and are particularly discernible in densely

structured substrata. Forma occulta has small, circular papillae exactly like C. vastifica. A "slight

spicular difference," that spiny microrhabds of C.

vastifica are "distinctly angulated, whereas those

of lampa are straight," was also pointed out by

Laubenfels (1950, 1953). The shape of this spicule

type, however, has been shown to vary considerably in C. vastifica, even within a single specimen

(Topsent, 1891), straight microrhabds being a

common type. C. corallinoides Hancock, characterized by flexuous microrhabds, is considered a mere

variation of C. vastifica (Topsent, 1891, 1900, 1932;

Volz, 1939). The formation of gemmules in C.

lampa, not reported by previous authors, is also

characteristic of C. vastifica (Topsent, 1932).
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

1i

FIGURE 19.—Cliona lampa, spicules: a, tylostyles; b, oxea; c, oxea and ipiny microrhabds.

(Magnification: a, b, X 520; c, X 1400.)

A worldwide revision of the Cliona vastifica

complex, which is beyond the scope of this paper,

will probably clarify also the status of the present

species. Only substantial revisionary data should

justify the elimination of a well-established name

like C. lampa.

The names chosen to designate the phenotypic

forms of Cliona lampa are derived from Latin,

occultus: hidden, concealed; flavidus: yellowish.

MATERIAL.—Forma lampa: USNM 24330; Station 4; 30 December 1969; in rock. USNM 24331;

Station 7; 6 January 1970; in pelecypod shell

(Chama). USNM 24332; Station 5; 12 January

1970; in dead coral (Diploria). Station 4; 30 December 1969; in rock. Station 5; 31 December

1969; in coral, pelecypod shell. Station 5; 30 December 1971; in pelecypod shell. Station 7; 6 January 1970; in pelecypod shell. Station 10; 11

January 1972. Station 12; 27 April 1971; in pelecypod shell. Station 21; 26 June 1972; in pelecypod

shell (Chama). Station 21; 12 January 1973.

Forma occulta: USNM 24339; Station 4; 26 December. 1969; in rock. USNM 24338; Station 9; 8

January 1970; in rock. USNM 24337; Station 5;

3 January 1972; in pelecypod shell. Station 2; 8

May 1971; in rock. Station 3; 26 December 1969;

in rock. Station 5; 31 December 1969; in pelecypod

shell. Station 5; 12 January 1970; in dead part

of live coral (Porites). Station 5; 30 December

1971; Station 7; 6 January 1970; in rock, pelecypod

shell. Station 8; 6 January 1970; in rock. Station

10; 11 January 1970; in rock, pelecypod shell. Station 11; 13 January 1970; in pelecypod shell, coral

rubble (Porites). Station 12; 27 April 1971; in

pelecypod shell. Station 13; 28 April 1971; in dead

coral. Station 17: 9 June 1972. Station 19; 7 June

1972; in rock. Station 20; 9 June 1972; in dead

coral (Montastrea). Station 21; 15 June 1972.

Forma flavida: USNM 24328; Station 10; 11

January 1972; in rock. USNM 24329; Station 21;

12 January 1973; in live pelecypod (Chama).

DISTRIBUTION.—Tropical western Atlantic.
NUMBER 165 25

FIGURE 20 Cliona lampa, spicules (SEM): Forma lampa: a, b, oxea with spines; c, spiny

microrhabds. Forma occulta: d, spiny microrhabds. Forma flavida: e, f, spiny microrhabds and

portion of oxeon. (Magnification: a, X 1300; b-e, X 4500; /, X 9000.)
26

Cliona amplicavata, new species

FICURES 21-23

DIAGNOSIS.—Papillae yellow, circular, discrete,

1.4 mm (mean diameter). Excavations typically

single, ovoid 10 X 7 mm (mean dimensions); filled

by yellow, soft, mucuous tissue. Tylostyles curved,

with spherical, mucronate heads, 262.5 X 7.1 um.

Rhaphides, in choanosome only, 129.8 X 0.8 um

(mean length X width).

DESCRIPTION.—Ectosome: The papillae of this

sponge are easily overlooked, although the single

large cavities containing the choanosome are most

conspicuous as soon as the substratum is split open.

The papillae are bright yolk yellow, circular, and

measure 0.8-2.6 mm (1.4 mm) in diameter. Some

bearing oscula can attain 3 mm with oscula of

0.5-1.0 mm. They occur in clusters of 2-5, but

they always remain separated. In undisturbed specimens they protrude 0.5-1.0 mm from the rock surface. Ostium papillae have the shape of a short

cylinder; oscular papillae are cone shaped. The

color in alcohol or when dry is dull gray. There

are no symbiotic algae. The most conspicuous histological feature is a number of large strongly

staining irregular bodies (Figure 21/), 35 X 24 um,

that contain calcareous fragments. These are possibly chips freed by the sponge during the burrowing process. A considerable amount of mucoid

material is contained in the exhalant canals.

Choanosome: This is yolk yellow like the ectosomal papillae. It is very soft, mucuous, and cavernous and fills the large chambers. In specimens

that have been kept outside the water for some

time, the tissue collapses and covers the chamber

walls only. The irregular bodies mentioned above

are also present here in abundance. In addition,

there are large, spherical, weakly staining cells without prominent inclusion. They measure 11-14 um

in diameter and contain a small anucleolate nucleus

(Figure 21e). Another very similar abundant cell

type contains basophilic granules of 1 um (Figure

21/). Choanocyte-chambers measure 30-35 um.

Excavations: One to five papillary perforations

(Figure 21a) serve each chamber. They are 0.8-

2.0 mm (1.3 mm) in diameter and connect with

the chamber by cylindrical or sometimes flaring

canals, 1.1-4.8 mm (3.2 mm) long. The chambers

(Figures 21c, d) are frequently single, ovoid, sometimes angular, and width and height are about

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

equal. The larger axis is usually parallel with the

substratum surface. The chambers measure 5 X

3 mm-22 X 12 mm (16 X 9 mm). Only few, if

any, adjacent chambers are communicating. In

that case they are separated by walls of 0.5-0.8 mm

thickness. In addition to natural crevices, there are

four to more than forty round or oval foramina

per chamber, 0.3-1.0 mm wide. Excavations reach

17 mm into the substratum.

Spicules: The tylostyles are usually curved (Figures 22a, b, 23«). In the papillae they are dense

and oriented perpendicular to the surface (Figure

21/;). This orientation is basically maintained also

in the choanosome, but the spicules are much less

common and many are placed in disorder. Their

heads are well set off, spherical, and typically raucronate; some ate distinctly subterminal. A dark

axial spot in the tenter of the head appears in

transmitted light In some specimens there is a

small percentage of styles. Bundles of fine rhaphides

(Figures 22b, 2$a-c) are common in the tissue of

the galleries. There are no spirasters or related

microscleres.

Spicule dimensions (in um): Tylostyles, length

X width: 190.0-290.0 X 4.5-8.0 (262.5 X 7.1).

Neck width: 3.2-6.4 (5.0). Head length X width:

4.8-12.0 X 4.8-10.4 (10.1 X 9.4). Rhaphides,

length X width: 117.5-150.0 X 0.8 (129.8 X 0.8).

REMARKS.—This species has the spiculation of

Cliona celata Grant. It differs from the latter by

the development of the papillae, by shape and size

of the burrow, by the consistency of the tissue, and

by the lack of incrusting and massive stages. No

rhaphides have been reported for American populations of C. celata (Hartman, 1958).

Cliona amplicavata can be confused with C.

flavifodina in the field. Its soft, mucuous tissue and

the more regular, ovoid chambers serve best for

characterization without looking at the spicules.

Siphonodictyon coralliphagum Riitzler, forma

obruta (Riitzler, 1971), another excavating sponge,

not yet known from Bermuda, resembles C. amplicavata by having a mucuous tissue and a very

similar burrow. In fact, from descriptions of the

sponge and from a sketch of the burrow (Noel P.

James, personal communication), I have previously, in absence of preserved material, erroneously

identified this sponge as Siphonodictyon. The

spiculation (oxea only) in the latter species, however, is very different.
NUMBER 165 27

V

•

•

••

•

*•• •

, 1

>

* t'

• .* .

Fir.iRE 21 Cliona amplicavata: a, papillary perforations; b, vertical section through papilla;

r, d, burrows with 1 and 2 papillary canals; e, choanocyte chamber and a-granular cells; /, granular cells and irregular bodies containing calcareous fragments. (Magnification: a, X 3; b, X 50;

c, d, X 3; e, f, X 850.)
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

O

FKUM 22—CKofM tmpltcmmt*, apkula: «, tyloatyto; *, tylottyle heads tad rhaphkk.

(Magnification: «, X 140; ft, X 580.)

1

•

a\

i

I

/ —"^ '

'1

FIGURE 23.—Cliona amplicavata, spicules (SEM): a, tylostyles and rhaphids; b, rhaphids; c,

magnified end of rhaphid. (Magnification: a, X 270; b, X 700; c, X 18000.)

The species name of Cliona amplicavata is derived from Latin amplus: large, spacious; cavatus:

hollowed out, and refers to the characteristic burrow.

MATERIAL.—USNM 24335 (holotype); Station

20; 9 June 1972; in dead portions of coral (Diploria). USNM 24333; Station II; 5 October 1969;

in rock. USNM 24334; Station 13; 9 May 1971; in

dead coral. USNM 24336; Station 17; 9 June 1972;

in dead portion of coral (Montastrea). Station 3;

26 December 1969; in dead coral.
NUMBER 165

Family SPIRASTRELLIDAE Ridley and Dendy

Spheciospongia othella Laubcnfels

FIGURES 24-26

Sphecioipongia othella Laubcnfels, 1950:94-96. fig. 42; pi. 2:

fig. 6.

Heterocliona cribaria.—Verrill, 1907:342-343. pi. 35D: figs. 2.3.

DESCRIPTION.—This species has a different appearance from the preceding clionids since, except

in very early stages, large parts of the sponge grow

on the surface of the rock. The epilithic portions

of the sponge consist of convex membranes, hollow pillows, hollow tubercular humps (Figure

24a), or smooth conical chimneys that cover or

protrude from vertical tunnels in the substratum

rock. Their diameters range from 3X 3 mm-30

X 20 mm; their height attains 1-30 mm. Only one

really large and still obviously burrowing specimen was found during the present survey. It was

irregular massive, 20 X 12 cm in horizontal dimensions, 5 cm in height.

Ectosome: This is represented by a deep black

or dark grayish brown outer layer, grayish black

in alcohol, dark gray when dry. The surface is

smooth; only near the base it is covered by adherent sediment particles. The oscula are 1-7 mm

in diameter (15 mm maximum in the large specimen). They have a raised collar, sometimes lighter

in color, and are strongly contractile. Ostia are

scattered over the surface and are less than 100 um

wide. A third kind of opening of undetermined

function is also abundant. These circular openings, also noted by Laubenfels (1950), stand in

groups, are not contractible, have no collars, and

measure 0.5-4 mm in diameter. A cross-section

through the wall of an epilithic chimney shows a

dense outer zone, about 100 um thick, of spheroid

or elongate cells (10 X 10 um-25 X 3 um), containing minute black pigment grains (Figure 24/).

These pigment cells occur scattered throughout

the tissue but, except for the outer layer, they are

only condensed where they form loose branching

strands. They are very rare in the innermost 100-

150 um zone, bordering the atrium. There they are

taken over by contractile cells. Granular cells, 16

X 16 um-30 X 10 um, containing small (1 um)

or larger (2 um) granules are abundant.

Choanosome: This is grayish to greenish ocher

29

and becomes dark gray in alcohol or when dry.

The fleshy tissue fills natural crevices and small

(new) tunnels; it thickly covers the walls of large

(old) tunnels (Figure 24 c). The central lumen

(exhalant canal) is circular in cross section and

measures 7-12 mm in diameter. Also the choanosome contains thin branching strands (50-80 urn

wide) of pigment cells. Choanocyte-chambers measure 25-30 um.

Excavations: There are essentially two size

classes of perforations produced at the substratum

surface (Figure 24 b). From the larger ones the

epilithic portions of the sponge protrude. There

can be one to several such structures, connected

by a common endolithic portion, which protrude

from holes 4-30 mm and more in diameter. In

advanced stages, the epilithic parts of a specimen

can be united to form a large massive sponge that

is still rooted in the substratum. The second,

smaller communicative perforations are probably

secondarily produced by outgrowth of the endolithic sponge mass. They measure 0.1-3.2 mm

(2.4 mm) in diameter, with a connecting canal

of 1-3 mm in length. The large openings lead

directly into flaring tunnels, which branch and

continue in horizontal and vertical direction under

the rock surface (Figure 24d). The tunnels are

circular or eliptical in cross section, ragged in

outline, and reach diameters of at least 22 X 10

mm. They are difficult to pursue throughout their

length. Tunnels can extend at least 100 mm into

the substratum.

Spicules: Tylostyles in the ectosome run in

strands parallel to the surface. Frequently these

strands branch and bend toward the surface, where

the spicules protrude in tufts (Figure 24c). Additional perpendicular tylostyles occur between the

tufts. Spirasters are rare and scattered. Tylostyles

in the choanosome are mostly scattered; some occur in strands. The shaft of the robust tylostyles

(Figure 25a, c) is quite uniform in thickness

through most of its length. The shaft is bent a

short distance behind the head. It does not taper

toward the point until approximately the last

fourth of its length. The heads are small, spheroid,

or elongate. Malformations are common. The

spirasters (Figures 256, c, 26) are small and ornate

with clusters of spines, particularly at both ends.

Spicule dimensions (in um): Tylostyles, length

X width: 115.7-373.8 X 1.5-8.9 (279.6 X 6.3).
I

I

FIGURE 24 Spheciospongia othella: a, pillow-shaped specimen protruding from substratum;

b, 2 size classes of perforations on substratum surface; c, tissue-covered tunnel inside rock;

d, cross section through 2 large tunnels, running below surface of coral rock; e, vertical section

through ectosome; /, pigment cells. (Magnification: a, b, d, X S; c, X 1; e, X 140; /, X 850.)
NUMBER ltt SI

a

FIGURE 25.—Spheciospongia othelta, spiculcs: a, tylostylcs; b, spiraster; c, spiraster and

tylosiyle head. (Magnification: a, X 140; b, c, X 1400.)

FIGURE 26 Spheciospongia othella, spicules (SEM): Spirasters. (Magnification: X 4500.)

Neck width: 1.5-8.1 (5.6). Head length X width:

3.7-14.8 X 3.7-11.5 (9.5 X 7.6). Spirasters, length

X width (shaft only): 8.0-25.6 X 0.8-3.2 (14.0 X

1.4). Spine clusters, maximum width: 2.8-5.6 (4.0).

Number of spines: 40-80+ (50+). Number of

bends: 1-5 (2.1).

REMARKS.—The preceding description is based

on specimens which were in an active burrowing

state. Large massive specimens, infrequently collected during the present survey, are described

by the original author (Laubenfels, 1950). It is

not established if and when, during the life cycle

of the species, the burrowing capacity stops.

MATERIAL.—USNM 24355; Station 2; 28 October

1969; in dead coral. USNM 24356; Station 10; 11

January 1970; in rock. USNM 24357; Station 13;

9 May 1971; in dead coral. Station 1; 26 October

1969; in dead coral. Station 10; 31 May 1972; in

rock. Station 11; 13 January 1970. Station 15; 3

May 1971; in rock. Station 17; 9 June 1972; in dead

coral. Station 19; 7 June 1972; in dead coral. Station 22; 26 June 1972; in rock.

DISTRIBUTION.—Bermuda.
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Literature Cited

Borojevil. R., W. G. Fry. W. C. Jones, C. L*vi. R. Rasmont,

M. Sari, and J. Vacelet

1968. Mise au point actucllc de la terminologic des

^sponges [A Reassessment of the Terminology for

Sponges]. Bulletin du Museum National d'Histoire

Naturelle, 39:1224-1235.

Carter, H. J.

1882. Some Sponges from the West Indies and Acapulco

in the Liverpool Free Museum Described, with

General and Classificatory Remarks. Annals and

Magazine of Natural History, 9(5):266-301. 346-

368, 2 plates.

Goreau, T. F., and W. D. Hartman

1963. Boring Sponges as Controlling Factors in the Formation and Maintenance of Coral Reefs. In R. F.

Sognnaes, editor, Mechanisms of Hard Tissue Destruction. American Association for the Advancement of Science Publication, 75:25-54, 16 figures.

Hancock, A.

1867. Note on the Excavating Sponges; with Descriptions

of Four New Species. Annals and Magazine of Natural History, 9(3) :229-242. 2 plates.

Hartman, W. D.

1958. Natural History of the Marine Sponges of Southern

New England. Peabody Museum of Natural History,

Yale University Bulletin, 12: 155 pages, 46 figures,

12 plates.

Hechtel, G. J.

1965. A Systematic Study of the Demospongiae of Port

Royal, Jamaica. Peabody Museum of Natural History, Yale University Bulletin, 20: 94 pages, 15 figures, 8 plates.

Laubenfels, M. W. de

1936. A Comparison of the Shallow-Water Sponges Near

the Pacific End of the Panama Canal with Those

at the Caribbean End. Proceedings of the United

States National Museum, 83:441-466, 6 figures.

1950. The Porifera of the Bermuda Archipelago. Transactions of the Zoological Society of London, 27: 154

pages, 65 figures, 2 plates.

1953. Sponges from the Gulf of Mexico. Bulletin of Marine Science of the Gulf and Caribbean, 2:511-557,

17 figures.

Neumann, A. C.

1966. Observations on Coastal Erosion in Bermuda and

Measurements of the Boring Rate of the Sponge,

Cliona lam pa. Limnology and Oceanography, 11:92-

108. II figures.

Pang. R. K.

In press. The Systematics of Some Jamaican Excavating

Sponges. Postilla.

RUtzler, K.

1971. Brcdin-Archbold-Smithsonian Biological Survey of

Dominica: Burrowing Sponges. Genus Siphonodictyon Bergquist, from the Caribbean. Smithsonian

Contributions to Zoology, 77: 37 pages. II figures,

9 plates.

In preparation. Participation of Burrowing Sponges in Erosion and Sediment Production on the Bermuda

Platform.

RUtzler. K.. and G. Rieger

1973. Sponge Burrowing: Fine Structure of Cliona lampa

Penetrating Calcareous Substrata. Marine Biology,

21:144-162, II figures.

Topsent. E.

1889. Quclqucs spongiaires du Bane de Campeche et de

la Pointe-A-Pltre. Mimoires de la Sociiti Zoologique

de France, 2:30-52, 12 figures.

1891. Deuxiemc contribution a l'ltude des Clionides. Archives de Zoologie Expirimentale et Generate, 9(2) :

555-592. 1 plate.

1900. £tude monographique des spongiaires de France,

III: Monaxonida (Hadromerina). Archives de ZooU

ogie Expirimentale et Ginirale, 8(3):331 pages. 8

plates.

1932. Notes sur des Clionides. Archives de Zoologie Expirimentale et Ginirale, 74:549-579, 11 figures.

Verrill, A. E.

1907. Porifera of the Bermuda Islands. Transactions of

the Connecticut Academy of Arts and Sciences, 12:

330-347. 2 plates.

Volz, P.

1939. Die Bohrschwamme (Clioniden) der Adria. Thalassia,

3: 64 pages, 16 figures, 5 plates.

Wells, H. W., M. J. Wells, and I. E. Gray

1960. Marine Sponges of North Carolina. Journal of the

Elisha Mitchell Scientific Society, 76:200-245. 63

figures.

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