Zoologica Scripta, Vol. 20, No. 3, pp. 195-199, 1991 0300-3256/91$3.00 + .OO Printed in Great Britain Pergamon Press plc 01991 The Norwegian Academy of Science and Letters

Coscinoderma sporadense sp.n. from the Aegean Sea with comments on Coscinoderma confragosum (Porifera, )

ELENI VOULTSIADOU-KOUKOURA, ROB W. M. VAN SOEST and ATHANASIOS KOUKOURAS

Accepted 11 November 1990

Voultsiadou-Koukoura, E., van Soest, R. W. M. & Koukouras, A. 1991. Coscinoderma spora- dense sp.n. from the Aegean Sea with comments on Coscinoderma confragosum (Porifera, Dictyoceratida) .-Zool. Scr. 20: 195-199.

Coscinoderma sporadense sp.n. is described from shallow waters in the North Aegean. This is the first record of a Coscznoderma species from the Mediterranean. The new species is compared to the deep water East Atlantic Coscinoderma confragosum Polkjaeff, 1884, and clear differences between them are established. The status of the genus Coscinoderma is discussed.

Eleni Voultsiadou-Koukouraand Athanasios Koukouras, Department of Zoology, University of Thessaloniki, 54006 Thessaloniki, Greece. Rob W. M. van Soest, Institute of Taxonomic Zoology (Zoological Museum), University of Amsterdam, P. 0. Box 4766, I009 A T Amsterdam, The Netherlands.

Introduction (BMNH 1888:8:9:1), which was kindly loaned to us by Miss S. M. K. Stone. The genus Coscinoderma Carter, 1883, as defined by Bergquist (1980), shows a disjunct distribution. Specifi- cally, it is found in shallow Australian waters [Coscino- Coscinoderma sporadense sp.n. (Figs 1-4) dermapesleonis (Lamark, 1814)] and in some deep water Type locality. Holotype and paratype (1) from Youra Island localities elsewhere, e.g. off Portugal (Coscinoderma (39"24'20"N, 24'8'45"E), North Sporades Islands, Aegean Sea, at a confragosum PolCjaeff, 1884) and off Florida (Coscino- depth of 3 m, on the sheltered surface of a rock. Paratype (2) from the derma lanuga de Laubenfels, 1936). Bergquist (1980) same area, in a dark cave located at a depth of 10 m. Paratypes (3) and (4) from Cape Drepanon (39"56'30"N, 23"57'20E), Sithonia Peninsula, refers most of the records of Coscinoderma, and espe- Aegean Sea, on the walls of a semidark cave at a depth of 15 m. cially those of von Lendenfeld (1889), to other genera. Type Material. Holotype Zoological Museum of Amsterdam (ZMA) Thus it may be concluded that species of Coscinoderma POR. 7700, collected 20.7.1984. Four paratypes (Ga 4488, Ga 4492, are few in number and generally of rare occurrence. collected20.7.1984; Ga4493, Ga4494, collected 13.7.1985), Museum of the Department of Zoology, University of Thessaloniki. Recent sampling in shallow waters (3-15 m) off the Etymology. The specific name is derived from the Sporades Islands, North Sporades Islands and off Cape Drepanon, both in the type locality. N. Aegean Sea, yielded several specimens of a spongiid , which could not be classified as one of the known Description Mediterranean keratose (Vacelet 1959; Pulitzer- Finali 1983; Pulitzer-Finali & Pronzato 1977, 1980). The The holotype (Fig. 1) is cushion shaped, with an average armoured surface and the largely non-anastomosing, thickness of about 5 mm and a surface area of 28 cm'. The entwined secondary fibers found in these specimens are two paratypes coming from the same geographical area not found in any species so far described from the Medi- have the same shape and thickness butt their surface area terranean. Instead, it was discovered that these charac- is much smaller. The other two paratypes are massive, ters answered to Bergquist's (1980) definition of Coscino- lobose, with a surface area of about 6 cm2 each. Colour is derma. The only description of a similar sponge remotely light brown becoming lighter in formaline. Both holotype in the same general biogeographic area is that of Coscino- and paratypes are soft, spongy and compressible. derma confragosum, given by PolCjaeff (1884). That The surface is conulose, the conules are about 1 mm species is only known from a single record from deep high and 2-4 mm apart. Oscules are few, 2-4 mm in waters (400 m) and it is ill-defined; Bergquist (1980) diameter. The ectosome is detachable. Perpendicular referred it doubtfully to Spongia. sections near the sponge surface showed that the ecto- It is the purpose of this paper to provide a description of some is armoured with sand grains and foreign spicules the Aegean Sea species and compare it to the only extant (Fig. 2a,b) and has a thickness of 100-350 urn. Qstia are specimen of Coscinoderma confragosum PolCjaeff, 1884 visible in some areas and have a diameter of 50-200 pm. preserved in the British Museum (Natural History) The ascending primary fibers (Fig. 3) have a diameter 195 Zoologica Scripta 20 196 E. Voultsiadou-Koukoura et al.

Fig. 1. Holotype of Coscinoderma sporadense sp.n. Scale in cm. of 50-80pm, and are cored with foreign material to such a spicules and sand grains, whitish-grey in colour, having degree, that sometimes spongin is hardly visible. The the form of a finely perforated plate when seen with the foreign material is usually sand grains mixed with low naked eye. The difference between this and the ectosome amounts of spicules, although some fibers are cored of C. sporadense (Fig. 2a) is that the latter, although also exclusively with spicules. The primary fibers are connec- made of sand grains and foreign spicules, lacks the reticu- ted to a dense, irregular, network of secondary fibers lated structure of the former. which in the vicinity of the primary fibers has the form of a As far as the skeleton is concerned, it was difficult to perforated plate (Fig. 3). As the distance from the pri- find primary fibers in C. confragosum and this may well be mary fibers and from the surface of the sponge increases, the reason that PolCjaeff makes no distinction between the secondary fibers become thinner and they hardly primary and secondary ones. However, the rare primary anastomose, but fork at intervals increasing with the fibers observed were cored with foreign material. The distance from the sponge surface (Fig. 4a). These inter- secondary fibers are uniform and form several broad vals are in most cases extremely long; they were not meshes. They fork at intervals of 100-400 pm [see also forking in intervals shorter than 1200 pm. Thus, the PolCjaeff (1884: 51, fig. l)].This is the main and, in our secondary network, in its greater part, resembles an opinion , most important difference between this species unwound clew (Fig. 4b). The secondary fibers have a and C. sporadense where the majority of secondary fibers diameter of 10-40 pm and they often terminate at a fork at intervals longer that 1200pm (Fig. 4). In addition, rounded or broadly acute end. the clear and dense reticulation of the secondary elements in the vicinity of the primary fibers of C. sporadense (Fig. Comparison with Coscinoderma confragosum Polijaefh 3) was not observed in C. confragosum where the second- 1884 ary fibers are very uniform in size and arrangement all over the sponge body. After examining the type material described by PolCjaeff Furthermore, the habitats of the specimens of C. con- (1884) we can make the following remarks. The original fragosum and C. sporadense differ in addition to the shape (massive, roundish, battledore shaped in longitudi- morphological differences. As mentioned in the Intro- nal section) was not recognisable since we examined only duction, C. confragosum was found off the coasts of a piece of the type specimen. However, the cribriform Portugal in a depth of 400 m, while our specimens were surface described by PolCjaeff is clearly visible; the easily found in shallow waters, 3-15 m, in the North Aegean detachable ectosome is a reticulation made of foreign Sea.

Zoologica Scripta 20 Coscinoderma sporadense sp. n. from the Aegean Sea 197

a

b

r

Fig .2. Ectosome of Coscinoderma sporadense sp.n.--a. Perpendicular section.-b. Tangcntial section

Discussion on the genus Coscinoderma emphasized the “cribriform incrustation” on the surface consisting of “quartz-grains” , and “fragments of sponge Despite Bergquist’s (1980: 458) opinion that C. confrago- spicules . . . as to constitute a . . . reticulate . . . structure, sum is “unrecognizable except as a very small fragment of whose interstices are uniformly subcircular”, and the a Spongia sp.”, we have demonstrated that the primary “Fibre . . . being almost uniformly alike , . . very small fibers are cored, the secondary elements are clear, fine and fine . . . scantily branched . . . very few are . . . found and numerous, that the surface is invested with a sand to contain foreign bodies.” Later, Carter (1885) described armour, but that the texture remains soft, spongy and two other specimens under the same name. compressible. These are the same characters Bergquist Von Lendenfeld (1889) gave the following definition: (1980: 457) mentioned in the description of Coscino- “Massive or flabellar Spongiidae with a very fine derma. Furthermore, the sponge body is massive, which is skeleton-net, a stout smooth sand cortex, and large con- one of the growth forms mentioned (Bergquist 1980: tinuous subdermal cavities without vestibular spaces.” He 457). Likewise, C. sporadense sp.n. answers in all its included C. lanuginosum, C. pyriforme Lendenfeld, 1889 aspects to Bergquist’s definition. What remains to be (which according to him had as synonyms the Caribbean established is whether Bergquist’s definition is really species Spongelia incerta and S. spinosa, both authored by based on Carter’s type species, i.e. C. lanuginosum Hyatt, 1875, but was apparently also recorded from South Carter, 1883. Australia), C. confragosum, C. polygonum von Lenden- Although Carter (1883) does not provide a generic feld, 1889 (from South Australia), and C. mathewsi (von definition, C. lanuginosurn is the only species mentioned Lendenfeld, 1886) (from the tropical Pacific and South in the original paper, so the description of that species Africa). may serve for the characters of the genus. Carter (1883) De Laubenfels (1948) considered Coscinoderma a Zoologica Scripta 20 198 E. Voultsiadou-Koukoura et al.

Fig. 3. Ascending primary fibers connected to the secondary network near the sponge surface.

Table I. A list of species allocated to Coscinoderma with comments on their status; bold printed names refer to valid species

C. altum Polejaeff, 1884. = Dysideopsis alta, according to Lendenfeld (1889); see also De Laubenfels (1948). C. concentricum Kirkpatrick, 1903. = Carteriospongia sp. according to Bergquist (1980). C. confrugosum Polejaeff, 1884. Occurrence: East Atlantic. C. densa (Hyatt, 1887) (see Whitelegge 1901) = Thorectaflabeliformis (Carter, 1885). C. denticulatum Polejaeff, 1884. = Spongia sp., according to De Laubenfels (1948). C. incerta (Hyatt, 1875). Junior synonym of C. pesleonis (Lamark, 1814), according to Bergquist (1980). C. lamarcki De Laubenfels, 1936, replacement name for Spongia sinuosa sensu Lamarck, 1814, referred to Coscinoderma by Topsent, 1930 (non: Spongia sinuosa Pallas, 1776). The redescription of Lamarck’s specimen by Topsent (1930) provides no indication that it is a Coscinoderma in the sense here employed; the specimen is considered a Hippospongia. C. Iunugu De Laubenfels, 1936. Occurrence: West Indies. C. lanuginosum Carter, 1883. Junior synonym of C. pesleonis (Lamark, 1814), according to Bergquist (1980). C. mathewsi Lendenfeld, 1889. = Spongia sp. according to Bergquist (1980). C. musicalis (Duchassaing & Michelotti, 1864). See Wiedenmayer (1977): table 48, and van Soest (1978). = Smenospongia aurea according to Kobluk & van Soest (1989). Phyllospongia myrobalana (Lamark, 1814), considered a senior synonym of Coscinoderma lanuginosum by De Laubenfels (1949), is in fact a Thorectid sponge according to Bergquist (1980). C.pesleonis (Lamark, 1814). Occurrence: South Australia, records by Lamark (1814), see Topsent (1930), Wiedenmayer (1989). C. polygonurn Lendenfeld, 1889. = Thorecta sp. according to Bergquist (1980). C. pyriforme Lendenfeld, 1889. Junior synonym of C. pesleonis (Lamark, 1814). C. sporudense sp.n. Occurrence: Eastern Mediterranean.

junior synonym of Phyllospongia and recognized apart from the clearly separatefoliascens, papyracea and vermi- Fig. 4. Two aspects of the secondary elements of Coscinoderma spora fera only pesleonis (Lamark, 1814), myrobalana dense n.sp., a being closer to the sponge surface than b. (Lamark, 1814) (with junior synonym Coscinoderma

Zoologica Scripta 20 Coscinoderma sporadense sp.n. from the Aegean Sea 199 lanuginosum) and lanuga De Laubenfels, 1936 as valid. Carter, H. J. 1883. Description of sponges from the neighbourhood of Port Philip Heads, South Australia.-Ann. Mug. nut. Hist 15: 301- C. confragosum was declared a “variety” of myrobalana. 321. Von Lendenfeld’s species were ignored by De Laubenfels De Laubenfels, M. W. 1936. A discussion of the sponge fauna of Dry (as was most of his work, except for a few plates and an Tortugas in particular and the West Indies in general, with material for a revision of the families and orders of the Porifera.-Pup. Tort. occasional remark). Lab. Carn. Inst. 30: 1-225. Bergquist (1980) distinguished sand-cortex bearing Dc Laubenfels, M. W. 1948. The order Keratosa of the phylum Phyllospongia, Carteriospongia and Coscinoderma as Porifera. A monographic study.-Occ. Pap. Allan Huncock Fdn. 3: 1-217. distinct, based on the shape, coring of the fibers and Kobluk. D. R. & Soest, R. W. M. van 1989. Cavity dwclling sponges in a the degree of development of the sand cortex and we southern Caribbean coral reef and their paleontological are in agreement with her analysis. In a later paper, implications.-Bull. Mar. Sci. 44: 1207-1235. Lcndenfeld, R. von 1889. A monograph of the Horny Sponges. Royal Bergquist et al. (1988) discussed Indo-Pacific Dictyocer- Society, London. atid genera with sand armour and non-anastomosing Polejaeff, N. 1884. Report on the Keratosa collected by H.M.S. Chal- secondary elements, and it was concluded that most of lenger during the years 1873-1876.-Rep. Challenger. Zool. 11: 1- 88. them fit the family rather than the family Pulitzer-Finali, G. 1983. A collection of Mediterranean Demospongiae Spongiidae; a notable exception was made of Coscino- (Porifera) with, in appendix, a list of the Demospongiae hitherto derma and in that, at least, we are in full agreement. recorded from the Mediterranean Sea.-Ann. Mus. Civ. Stn. nut. Genova 84: 445-621. Most features of C. sporadense resemble those of the Pulitzer-Finali, G. & Pronzato, R. 1977. Report on a collection of bath sponge genera Spongia and Hippospongia, except Sponges from the Bay of Naples. 11. Keratosa. Pubbl. Stuz. zool. for the peculiar unbranched secondary fibers and the Nupoli 40: 83-104. Pulitzer-Finali, G. & Pronzato, R. 1980. The Keratosa in a collection of surface armour. Mediterranean Sponges mainly from the Italian coasts.-Ann. Mus. All specimens referred to or allocated to Coscinoderma Civ. Stn. nut. Genova 83: 127-158. in the literature are listed in Table 1. Soest, R. W. M. van 1978. Marine sponges from Curacao and other Caribbean localities. Part I. Keratosa.-Stud. Fauna Curacao Curibb. Id. 56: 1-94. Topsent, E. 1930. Eponges dc Lamarck conservkes au Museum de Paris. Deuxitme partie.-Arch. Mus. Nut. Hist. nut. 6: 1-60. References Vacelet, J. 1959. Rkpartition generale des Cponges et systCmatique des eponges cornees de la region de Marseille et de quelques stations m6diterraneenes.-Rec. Trav. Stn. mar. Endoume 26: 39-101. Bergquist, P. R. 1980. A revision of the supraspecific classification of the Wiedenmayer, F. 1977. Shallow-watcr sponges of the Western orders Dictyoceratida, Dendroceratida and Verongida (class Bahamas.-Experientiu Supplementu 28: 1-287. Demospongiae).-N. Z. J. 2001. 7: 443-503. Wiedenmayer, F. 1989. Demospongiac (Porifera) from Northern Bass Bergquist, P. R., Ayling, A. M. & Wilkinson, C. R. 1988. Foliose Strait (shelf of Southern Australia).-Mem. Mus. Vicroria 50: 1-242. Dietyoceratida of the Australian Great Barrier Reef. 1. Whitelegge, T. 1901. Report on the sponges from the coastal bcaches of and phylogenetic relationships.-Mar. Ecol. 9: 291-319. New South Wales.-Rec. Austral. Mus. 4: 55-118.

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