Bijdragen tot de Dierkunde, 54 (2): 211-219—1984

Deficient Merlia normani Kirkpatrick, 1908, from the Curaçao

reefs, with a discussion on the phylogenetic interpretation

of sclerosponges

by

R.W.M. van Soest

Institute of Taxonomie Zoology (Zoölogisch Museum), University of Amsterdam,

P.O. Box 20125, 1000 HC Amsterdam, The Netherlands

Summary INTRODUCTION

basal A specimen of the sclerosponge Merlia lacking a skeleton of calcareous chambers (corresponding to the Kirkpatrick (1908) described Merlia normani description of Merlia deficiens Vacelet, 1980) is reported from Porto Santo, Madeira Archipelago. This from the underside of the coral of 10 Agaricia at a depth m basal skeleton of calcareous species possesses a on the reef of Curaçao, Netherlands Antilles. Its spicula- chambers, in which and over which the soft tion is identical to Merlia normani Kirkpatrick, 1908 of the These soft parts are draped. parts specimens with the basal skeleton, from which it is con-

include a mineral skeleton of characteristic cluded that M. deficiens and M. normani are probably con- SÌO2 specific. spicules: tylostyles, rhaphides, commata and

The phylogenetic significance of a sclerosponge with an clavidiscs. Subsequently, the species has been unstable calcareous skeleton is discussed and in accor- found in the Indian Ocean (Dendy, 1922), the dance with Vacelet (in press) it is postulated that Mediterranean (e.g. Vacelet, 1980; Pulitzer- sclerosponges are polyphyletic. It is suggested that

skeletons ancestral character for and & sclerosponge are an a Finali, 1972) Jamaica (Hartman Goreau,

of the The of large part Demospongiae. relationships 1970).

Demospongiae and Calcarea are discussed in the same Pouliquen (1972) was the first to record light. Merlia without its characteristic skeleton (from a

cave near Marseille); Vacelet (1980) compared

the forms and concluded that were Résumé two they closely related, but probably specifically distinct Un du Merlia dépourvu du exemplaire Sclérospongiaire of clavidiscs in on account consistently larger squelette basal de logettes calcaires, et correspondant à ce the deficient form, which he named Merlia defi- qu’avait été décrit comme Merlia deficiens Vacelet, 1980, a ciens. été découvert à face inférieure corail à 10 la du Agaricia m It will be shown below that these differences de profondeur sur le récif de Curaçao, Antilles Néerlan- daises. La est à celle des and that Vacelet' spiculation identique exemplaires are slight s new species is à squelette basal de Merlia normani Kirkpatrick, 1908, d’où doubtful. Thus it appears that within one and il est possible de conclure à la conspécificité probable de the the calcareous base be same species may M. deficiens et de M. normani.

facultative or unstable. The On discute de la signification phylogénétiqued’un Sclé- phylogenetic

à calcaire instable d’accord of this fact is discussed below in the rospongiaire squelette et, avec significance

Vacelet (sous presse) on conclut au caractère polyphyléti- of Vacelet's ideas dur- light newest put forward des On les que Sclérospongiaires. suggère que squelettes ing a recent symposium (September, 1983) des Sclérospongiaires représentent un caractère ancestral in For the soft (Vacelet, press). this reason parts des Dans le même pour une grande partie Demospongiae. of various other have ordre discute des relations de sclerosponges been d’idées, on parenté entre well. Demospongiae et Calcarea. studied as

Downloaded from Brill.com10/04/2021 05:18:18AM via free access 212 R. W. M. VAN SOEST - DEFICIENT MERLIA

Figs. 1-4: 1, Clavidisc of deficient Merlia normani from the Curaçao reefs. 2, Head of tylostyle of deficient Merlia normani

3, Acanthose comma of deficient Merlia normani. 4, Overall view of spicule complement of deficient Merlia normani, show- ing clavidisc, tylostyle and acanthose rhaphides.

SYSTEMATIC DESCRIPTION extreme thinness, but from scrapings it is clear

that whispy bundles of tylostyles form the main Order POECILOSCLERIDA support; microscleres are numerous. Family Biemnidae Spicules: tylostyles with sinuous shaft and

small but distinct heads: 137-160 1-2 by [xm; Merlia normani Kirkpatrick, 1908 clavidiscs (compound diancistras): 42-57 by

Merlia normani Kirkpatrick, 1908: 510, pl. XV 10-18. 21-34 in thick figs. [im; rugose (acanthose) rhaphides Merlia deficiens Vacelet, 1980: 227, pl. I figs. 3-4; Vacelet, dragmata: 44-84 by 0.5 fxm; sigmatose com- 1981: 321. 7-15 mata: (Jim.

Material. — Zoologisch Museum Amsterdam coll. no.

ZMA POR. 4th W. of 4765, Curaçao, buoy (500 m

10 surface of Piscadera), depth m, under Agaricia colony, DISCUSSION 2-1-1981, coll. L. Delvoye.

Description (figs. 1-5). — Extremely thin (~ The decision to unite normani and deficiens is

0.5 about 2 in made mm), incrusting patch 2 cm on the basis of spicule comparisons of lateral expansion; in alcohol the specimen is in- available Merlia material and literature data (cf. visible the of the coral that is dif- against background table I). Vacelet's suggestion there a substrate. In life the colour was red. The ference in clavidisc size between normal and skeletal is difficult due the deficient structure to study to specimens seems to be correct, but the

Downloaded from Brill.com10/04/2021 05:18:18AM via free access 54 - 1984 213 BIJDRAGEN TOT DE DIERKUNDE, (2)

TABLE I

Spicule sizes (μm) of Merlia specimens.

Locality Habit Authority Tylostyles Clavidiscs Rhaphides Commata

144-152/2-3 30-44.0-53/29-36 48-74 14-15 Porto Santo, sclerosponge present paper

Madeira Archipelago

Mediterranean sclerosponge Vacelet, 1980 — 38.6 — —

Mediterranean sclerosponge Pulitzer-Finali, 1972 136-153/1.5 41-47 54-80 15-17

Jamaica sclerosponge Hartman & Goreau, 142-171 28-33/24-28 — —

1970

Jamaica sclerosponge Vacelet, 1980 — 34.3 — —

Mediterranean deficient Vacelet, 1980 — 56.7 — —

Curaçao deficient 137-160/1-2 42-49.3-57/21-34 44-84 7.5-15 present paper

Mediterranean unknown Pulitzer-Finali, 1983 122-173/1.5-2 29-78 44-90 —

Chagos Archipelago, unknown Dendy, 1922 — 57-66/41 — —

Indian Ocean

is matching of the other spicule sizes so great structural correspondence between the dian-

that A cistras of Hamacantha and the clavidiscs of a separate specific status seems doubtful.

further for the is found in Merlia. These characters be reason uniting two shared can only ex-

the geographic distribution of both forms, plained in one way: shared ancestry at the fami-

which overlaps completely in the tropics and ly level.

subtropics.

The acceptance of this has the heavy implica- PHYLOGENETIC IMPLICATIONS OF THE tion that the calcareous basal skeleton is SCLEROSPONGE SKELETON

unstable within the species, a suggestion from Merlia normani I able Apart was to study forward Vacelet already put by (1980). specimens of several other sclerosponges (listed The polyphyly of the "class" Sclerospongiae in the Acknowledgements below), which allows will be discussed below. Vacelet (1979) erected me to draw firsthand conclusions. I did not look order Merliida for Merlia within the a separate at fossil specimens, but discussions with Mr. Demospongiae, which was followed by Pulitzer- Theo van Kempen (Geological Institute, Finali (1983). For various reasons this is not ac- of and Mr. Theo University Amsterdam) van cepted here, the most important being the great Koolwijk, who did a literature survey of fossil correspondence in spiculation of Merlia with hypercalcified , contributed in shaping members of the poecilosclerid family Biem- my opinion. nidae. In a previous paper (Van Soest, 1984) I outlinedthis others family as comprising among The polyphyletic nature of the class Sclerospongiae the genera Biemna Gray (1867), Desmacella

Schmidt (1870), Hamacantha Schmidt (1870), The existence of this class is widely accepted and Hechtel Merlia normani Neofibularia (1965). among neontologists (e.g. Hartman & Goreau,

shares the tylostylote megascleres with 1970, 1975; Hartman, 1972, 1979; Levi, 1973;

the microxea and in The Desmacella, roughened (rhaphides) Bergquist, 1978, press). only con-

with the with Biemna and fessed critic Neofibularia, commata among recent spongologists is

and there is Vacelet and in Neofibularia, finally an undeniable (1979, 1980, 1981, press), and

Downloaded from Brill.com10/04/2021 05:18:18AM via free access 214 R. W. M. VAN SOEST - DEFICIENT MERLIA

spirasters; this combination is characteristic of

the genus Spirastrella, family Spirastrellidae,

order Hadromerida.

Calcifibrospongia:

No material has been studied, but from Hart-

man's (1979) well-presented description it can

be that the easily seen strongyle reticulation

found in this form shows undeniablesimilarities

with haplosclerids of the family Haliclonidae.

The data above 6. given are summarized in fig.

If we should wish to unite these forms into a

single (sub-)class, comprising sclerosponges on-

admit: ly, we are forced to

— occurred either that there has an improbable

number of parallel developments in spicule

categories and combinations in Sclero-

the hand and Demo- spongiae on one

the spongiae on other,

— or that all these spicules are primitive,' occur-

ring already in the (Cambrian?) common Line of of deficient Fig. 5. drawings spicules Merlia nor- of both ancestor groups. mani. As several of these spicule types are quite

unusual limited such with only a distribution,

his is here A brief reasoning supported. survey conclusions are hardly parsimonious. For this

of the is show I Vacelet in his recent sclerosponges necessary to reason strongly support refuta-

the of the evidence the existence weight against tion of the Sclerospongiae as a class separate

of such found in the soft of these a class, parts from the Demospongiae.

sponges.

The next question to be solved is: has the basal

Ceratoporella, Goreauiella, Stromatospongia, Hispido- calcareous skeleton been developed separately

and petra Astrosclera: in each different line, or is it a primitive

These all share viz. of a single spicule type, an (ancestral) character of a large group

with the thorns in does acanthostyle arranged ver- orders? Merlia not provide us

ticils. Such characteristic is also with If a spicule type an unequivocal answer to this. parallel

found in is a single non-sclerosponge genus: development postulated, one might cite

This of Merlia form in which Agelas. "enigmatic" group species was as an example of a living

raised has recently to ordinal status, but the development of a basal skeleton is actually

axinellid affinities. Astrosclera is witnessed this in time. presumed very moment Adversely,

known to have unstable if it is assumed in has spicules, as specimens (as Vacelet, press, put

have been found with without them or only a forward) that the basal skeleton is primitive,

few This fact is (Vacelet, 1977, 1981). given shared with many fossil sponges (stromatopo-

considerable in be weight arguments presented roids, chaetetids), then Merlia may cited as

below. of in the of an example a process loosing

its calcareous base. However, three more

Acanthochaetetes = for the view ( Tabulospongia): arguments latter can be put for-

in the soft and The spicules parts are tylostyles ward: firstly, the fossil record of sclerosponges

Downloaded from Brill.com10/04/2021 05:18:18AM via free access - 215 BIJDRAGEN TOT DE DIERKUNDE, 54 (2) 1984

6. Distribution of the orders redrawn from Fig. sclerosponge genera over currently recognized demosponge (partly Berg-

quist, 1978, fig. 8.8).

is quite extensive (the sclerosponge nature of would be conceivable that they evolved from

chaetetid tabulates the calcareous many stromatoporoids and stromatoporoids by replacing

chamber walls it seems to be firmly established, see e.g. Stel, by SÌO2 spicules). Adversely,

1978, cf. also Hartman et al., 1980); secondly, could be argued that the earliest (Middle Cam-

is doubtful the number of living sclerosponges quite low, brian) stromatoporoids are sponges

low of the known pointing to a strategic advantage (Stearn, 1979), unequivocal ones being

basal calcareous which in its from the Middle Ordovician. This would skeleton, turn sup- only

the idea of it been abandoned that the lithistids could be ports having by mean considerably

for and have been ancestral to the most sponges competitive reasons; thirdly, older, may

the cryptic habitat of living sclerosponges points stromatoporoids. We will leave this question

in the direction: it is that undecided here. same probable they

have been into unfavourable niches Lithistids have limited distribution pushed by a among

modern reef building organisms (stony corals the Demospongiae, being confined mostly to

and non-sclerosponges). Vacelet's (in press) astrophorid and spirophorid tetractinomorphs,

overall thus with the few unrelated sublithistids picture is consistent a probably excepted.

Those forms considered incertae Lévi observed facts and represents the most accept- sedis by

of the lack of able hypothesis. (1973) on account true tetracts

lost these and quite likely just spicules are prob-

Some remarks lithistid in related. as the on sponges are ably closely Just sclerosponges,

order here. lithistids assume the the lithistids have been likewise driven In many ways to an un-

habit and it is that same as sclerosponges, likely favourable habitat (they are commonest in deep

of they represent a separate attempt sponges to water beyond the reefs), due no doubt to com-

the reef habitat. From the literature occupy petition with more successful reefbuilders.

(e.g. Finks, 1970, cited in Hartman et al.,

1980: 74) it could be concluded that lithistids SPHINCTOZOANS, MURRAYONIDS appeared (in geological time) after the

and could with stromatoporoids, thus not cer- These are "living fossils" described or

be considered redescribed Vacelet tainty a parallel development (it recently by (1970, 1977,

Downloaded from Brill.com10/04/2021 05:18:18AM via free access 216 R. W. M. VAN SOEST - DEFICIENT MERLIA

the class Vacelet 1979), assigned or formerly assigned to probably stromatoporoids. Although

Calcarea. This class is well characterized by a (1979) has shown that the only living sphincto-

derived has number of synapomorphies (shared zoan, Vaceletia crypta, Demospongiae

the of which is the characters and the characters), most important lacks Calcarea synapomor-

possession of triactine and tetractine CaCC>3 phies, nevertheless comparable fossil forms with

been spicules. The origin of the group has never typical Calcarea spicules have been described,

satisfactorily clarified, because the fossil record which have to be attributed to the Calcarea,

is and the shows few Verticillites. poor group plesiomor- e.g.

phous characters. Contrary to the opinion of Borojévic (1979),

The sclerosponge-like murrayonids give food these data and observations may be interpreted

such that the of the Calcarea to the assumption that such a synplesiomorphy in a way origin

of Calcarea other be found in from and sponges may early demosponge-stromatoporoid-sphinc-

made series their common stromatoporoid origin, an idea tozoan stock is plausible through a

advocated by Termier & Termier (1973). If this of presumed steps: First, it is suggested that

evolved is rejected (as Vacelet, in press, apparently has primitive Sphinctozoa-like sponges

basal from their siliceous done), then we have to assume that a stromatoporoids by loosing

calcareous skeleton has at least evolved twice in- spicules. The tendency to loose the siliceous

from soft-bodied is is observed in the dependently sponges (this spicules recent Astrosclera;

not entirely impossible, since there are quite most other demosponge-sclerosponges have

large structural differences between different feebly developed spicules compared to their

sclerosponge basal skeletons). non-sclerosponge relatives (compare spicule

Where has the bodied sizes of Merlia with those of dialytine (soft sponges most biemnids,

with free calcareous spicules) structure come those of Acanthochaetetes with spirastrellids).

from? Until this of Hereafter loss of basal recently, type sponge was a progressive the

known from the Carboniferous onwards calcareous skeleton occurred, unrelated to the

cited in Hartman et 1980: loss of in the it is assumed (Wendt, 1979, al., spicules, same way as

170), but Van Kempen (1978) rather casually to have happened in demosponge sclerospon-

mentioned the of and indeed is witnessed in Merlia and in the occurrence dialytine spicules ges,

from the Middle Cambrian of Australia. murrayonids Murrayona and Paramurrayona.

careful and total loss of However, after restudy reconsidera- Finally, the resulting a supporting

would in have been tion (photographs and comparisons with recent skeleton some cases com-

Van and I reached the of of dialytine sponges), Kempen pensated by development a new type the conclusion that the the spicules are not likely to support, calcareous triactine or tetractine

in in form similar that of certain be dialytine origin. They probably represent spicules, a to tetractinellid In is spicules. a forthcoming paper demosponge SÌO2 spicules (calthrops). It con- of Van the of these ceivable that the of Kempen presence particular ability making SÌO2 spicules forms of will be discussed in lost the and that the of spicules more was along way ability detail. Consequently, it still stands that soft- making a certain type of spicule was retained. bodied calcareous out of the sponges came blue,

HOMOSCLEROPHORIDA somewhere near the beginning of the Mesozoic, in which time and inozoans sphinctozoans were Dialytine Calcarea show similarities with already well established (Wendt, 1979, cited in , which have always been Hartman The earliest et al., 1980). sphincto- considered but it independently acquired*, may in the zoans apparently developed Middle

* Möhn in a recent major revision of the classifica- it is (1984) Cambrian (Picket & Jell, 1983), so impossi- tion of the Porifera, apparently considers these similarities ble to any other suggest evolutionary sequence them be as primitive, since he clearly states to homologous than to out that in the same in both but the time he considers Calcarea point they appeared groups, at same

tetractinellids and the oldest and period as lithistids, (?) as most original sponge group.

Downloaded from Brill.com10/04/2021 05:18:18AM via free access BIJDRAGEN TOT DE DIERKUNDE, 54 (2) - 1984 217

7. overall of A: view with Fig. Diagrams representing phylogenetic relationships major sponge taxa. Currently accepted

classes Hexactinellida, Calcarea and Demospongiae (Tetractinomorpha + Ceractinomorpha + Homoscleromorpha).

B: suggested view based on the idea that hypercalcified skeletons were developedonly once and on the assumption that

similarities Calcarea of and Homoscleromorpha are homologous.

be worth wile to reconsider this. Some of these The amphiblastula found in Calcarea

similarities form and and is similar in are: calthrops-like spicule Homosclerophorida structure

amphiblastula larva. Donadey (1979) noted and function, but according to Tuzet (1948) this

several be considered corresponding cytological phenomena, must as a parallel development

be because the in which might well turn out to genuine embryology both cases differs

the I homologies. Possibly, large choanocyte substantially. am not certain of the weight of chambers in both be these differences. The similarities and limited groups may a homologous

distribution feature as well. among sponges as a whole make

the of hard: The calthrops-like spicule form was con- assumption a parallel development

it too muchof coincidence. The sidered primitive by many older spongologists seems a simpler

of larva because of its similarity to large tetractine type found in Clathrina is quite naturally

reduction of the spicules as orthotriaenes, and to astrose explained as a amphiblastula.

A further of microscleres found in tetractinellids and maybe corroboration of shared ancestry

Calcarea and be also in hexactinellids. The more recent view Homosclerophorida may

the found in the fact that in in- emphasizes monaxone spicule as primitive, Homosclerophorida because of its wider distribution deed known without among sponges some sponges are any

skeleton at all These here in general. This matter is left here undecided, (e.g. Oscarella). are

assumed be for the the to a in favour of the except suggestion that calthrops may testimony be for Calcarea and Homo- that but unrelated a synapomorphy hypothesis two successive,

reduction from sclerophorida. processes some stromatoporoid-

sphinctozoan stock to the naked stage have in- in his the oldest line Homosclerophorida, view, represent deed occurred at a certain moment in the evolu- in the Demospongiae. This view clearly violates the princi- tion of these assorted of groups sponges. Ver- of because both small in diversi- ple parsimony, groups are and ticillites, Vaceletia, murrayonids are ex- ty, with few taxa, and without extensive fossil record. Next

there smaller and inconsistencies of relicts of that have to this, are many larger amples lines diverged in while number of Möhn's new the dazzling new- from this system, evolutionary sequence. ly proposed subkingdoms, phyla, subphyla and classes The dissimilarities many and perhaps the seems unprecedented. The discussion of these proposals, doubtful homology of some of the similarities however, is the of the which is beyond scope present paper, contribute the ofthe together to that intended only to explore some possible consequences assumption any of the of Calcarea acceptance of the polyphyly of the Sclerospongiae. common ancestor and Homo-

Downloaded from Brill.com10/04/2021 05:18:18AM via free access 218 - R. W. M. VAN SOEST DEFICIENT MERLIA

there sclerophorida (if has been one at all) lived Trans. Linn. Soc. London, (2, Zool.) 18: 1-164,

I-XVIII. at least as far back as Devonian times. If the pis. DONADEY, O., 1979. Contribution à l'étude cytologique would be present suggestions eventually de deux Homosclerophoridés: Oscarella lobularis substantiated by new corroborating evidence (Schmidt) et Plakina trilopha Schulze. In: C. LÉVI

of new (reexamination embryology, fossils, & N. BOURY-ESNAULT eds., Biologie des Spongiaires.

revision of the its Cent. Homosclerophorida), conse- Colloques int. natn. Rech, scient., 291:

165-172. would be that the Calcarea would have quence HARTMAN, W. D., 1972. Ceratoporella (Porifera: Sclero- in the to be incorporated the Porifera system at spongiae) and the chaetetid "corals". Trans. Conn.

same level as the Homosclerophorida (at pres- Acad. Arts Sci., 44: 133-148.

ent: In view of the subclass). increasing 1979. A new from the Bahamas and , sclerosponge

evidence of calcareous and siliceous skeletal its relationship to Mesozoic stromatoporoids. In:

C. LEVI N. BOURY-ESNAULT material in the & eds., Biologie des being employed same sponge, Spongiaires. Colloques int. Cent. natn. Rech, this seems acceptable. A tentative diagram of scient., 291: 467-474. the presumed phylogenetic relationships is HARTMAN, W. D. & T. F. GOREAU, 1970. Jamaican in 7B. coralline their and presented fig. sponges: morphology, ecology

fossil relatives. Symp. zool. Soc. London, 25:

ACKNOWLEDGEMENTS 205-243.

& 1975. A Pacific tabulate , sponge, living Dr. Jean Vacelet (Marseille) and Prof. Dr. Patricia Berg- of of representative a new order sclerosponges. Postil- quist (Auckland) kindly presented me copies of their la, 167: 1-21. respective lectures given at the London symposium, HARTMAN, W. D., J. WENDT & F. WIEDENMAYER, 1980. September 1983. Drs. Laurent Delvoye collected and and fossil 8: 1-274 Living sponges. Sedimenta, donated the specimen of Merlia normani from the Curaçao (University of Miami, Florida). reef. Dr. Rolf Bak donated (Texel) a specimen of TH. M. G. 1978. KEMPEN, VAN, Anthaspidellid sponges Goreauiella auriculata from Jamaica (Zoölogisch Museum from the Early Paleozoic of Europe and Australia. Amsterdam coll. no. ZMA POR. 4798), Drs. Eric Newton Neues Jahrb. Geol. Paläont. Abhandl., 156: 305- donated of (Bonaire) a specimen Stromatospongia norae from 337. Jamaica (ZMA POR. 4572), Dr. Steven Weinberg (Lux- 1908. On of KIRKPATRICK, R., two new genera recent embourg) donated a specimen of Ceratoporella nicholsoni Ann. pharetronid sponges. Mag. nat. Hist., (8) from Jamaica (ZMA POR. 4569), Dr. Maya Best 2: 503-515, pis. XIII-XV. (Leiden) donated a specimen ofAcanthochaetetes wellsi from LEVI, C., 1973. Systématique de la classe de Demospongi- the Philippines (ZMA POR. 5323), Dr. Harry ten Hove aria (Démosponges). In: P. P. GRASSE ed., Traité (Utrecht) and Dr. Helmut Zibrowius (Marseille) donated de zoologie. Anatomie, systématique, biologie. specimens of Petrobiona massiliana from Marseille (ZMA Tome 3, fascicule 1. Spongiaires: 577-631 (Masson, POR. 4467, 5324). Miss S. M. K. Stone (British Museum Paris). (Natural History), London) and Drs. J. C. den Hartog MÖHN, E., 1984. System und Phylogenie der Lebewesen. (Rijksmuseum van Natuurlijke Historie, Leiden) con- Band 1: i-xxxv, 1-884, pis. 1-318 (Schweizerbart, sented in the loan of Merlia normani specimens from Porto Stuttgart). Santo. Drs. Theo of van Koolwijk (Institute Taxonomie PICKET, J. P. & P. A. JELL, 1983. Middle Cambrian of is thanked for his Zoology, University Amsterdam) con- Sphinctozoa (Porifera) from New South Wales. structive comments. Mem. Ass. Australas. Palaeontols., 1: 85-92.

POULIQUEN, L., 1972. Les Spongiaires des grottes sous- REFERENCES marines de la région de Marseille. Téthys, 3:

BERGQUIST, P. R., 1978. Sponges: 1-268, pis. 1-10 717-758.

(Hutchinson University Library, London). PULITZER-FINALI, G., 1972. Report on a collection of

in Poriferan Proc. Ass. from the of I. , press. relationships. Syst. sponges Bay Naples, Sclerospongiae,

Symposium "On origin and relationships of lower Lithistida, Tetractinellida, Epipolasida. Pubbl. Staz.

invertebrates", London, Sept. 1983. zool. Napoli, 38 ("1970"): 328-354.

1979. Evolution des Calcarea. In: C. LEVI 1983. A collection of Mediterranean BOROJÉVIÖ , R., , Demospon-

& N. BOURY-ESNAULT eds., Biologie des Spongiaires. giae (Porifera) with, in appendix, a list of the

Cent. Colloques int. natn. Rech, scient., 291: 527- Demospongiae hitherto recorded from the Mediter-

530. ranean Sea. Annali Mus. civ. Stor. nat. Giacomo

DENDY, A., 1922. Report on the Sigmatotetraxonida col- Doria, 84: 445-621.

lected H.M.S. "Sealark" in the Indian Ocean. R. W. M. 1984. Marine from by SOEST, VAN, sponges

Downloaded from Brill.com10/04/2021 05:18:18AM via free access BIJDRAGEN TOT DE DIERKUNDE, 54 (2) - 1984 219

and other Caribbean localities. Part III. de française, de Curaçao sponges Polynésie Madagascar et

Poecilosclerida. Stud. Fauna Curaçao, 199: 1-167, de La Réunion. Bull. Mus. natn. Hist. nat. Paris,

pis. I-X. (3) 444 (Zool. 307): 345-368.

C. W., 1979. and related 1979. et affinités d'une Éponge STEARN, Stromatoporoids , Description

organisms. Fossil int. Newsletter, 8 (1): Sphinctozoaire actuelle. In: C. LÉVI & N. BOURY- 49-50. ESNAULT eds., Biologie des Spongiaires. Colloques

STEL, J. H., 1978. Studies on the paleobiology of int. Cent. natn. Rech, scient., 291: 483-493.

favositids: 1-247 of Gro- 1980. calcaire facultatif et de (Ph.D. Thesis, University , Squelette corps

dans le ningen). régénération genre Merlia, Eponges

TERMIER, H. & G. TERMIER, 1973. Stromatopores, apparentées aux Chaetétidés fossiles. C. R. Acad.

Sclérosponges et Pharétrones: Les Ischyrospongia. Sci. Paris, (D) 290: 227-230, pl. I.

Annls. Mines 26: 285-297. 1981. Éponges Géol., , hypercalcifiées ("Pharétronides",

TUZET, O., 1948. Les premiers stades du développe- "Sclérosponges") des cavités des récifs coralliens

ment de Leucosolenia botryoides Ellis et Solander de Nouvelle-Calédonie. Bull. Mus. natn. Hist,

et de Clathrina (Leucosolenia) coriacea Mont. Annls. nat. Paris, (4) 3 (A, 2): 313-351.

Sci. 11: 103-113. in Coralline and the evolution of nat., (Zool.) , press. sponges

VACELET, J., 1970. Les Éponges Pharétronides actuelles. the Porifera. Proc. Syst. Ass. Symposium "On origin

Symp. zool. Soc. London, 25: 189-204. and relationships of lower invertebrates", London,

1977. Éponges Pharétronides actuelles et Scléro- 1983. , Sept.

Received: 6 July 1984

Downloaded from Brill.com10/04/2021 05:18:18AM via free access