Sponges, Genus Mycale (Poecilosclerida: Demospongiae: Porifera), from a Caribbean Mangrove and Comments on Subgeneric Classification
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23 December 1998 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):737-773. 1998. Sponges, genus Mycale (Poecilosclerida: Demospongiae: Porifera), from a Caribbean mangrove and comments on subgeneric classification Eduardo Hajdu and Klaus Riitzler (EH) Institute of Systematics and Population Biology, University of Amsterdam, P.O. Box 94766 1090-GT, Amsterdam, The Netherlands; Present address: Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, sin 20940-040, Rio de Janeiro, RJ; and Centro de Biologia Matinha, Universidade de Sao Paulo, Sao Sebastiao, Sp' Brazil; (KR) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.-Eight species of Mycale Gray (Mycalidae, Poecilosclerida, De mospongiae) are described from marine mangroves on the barrier reef of Be lize, Central American Caribbean. Two are new: Mycale (Aegogropila?) car migropila and M. (Ae.) citrina. Other species found are M. (Ae.) amdti, M. (Arenochalina) laxissima, M. (Carmia) magnirhaphidijera, M. (C.) microsig matosa, M. (Mycale) laevis and M. (Paresperella) sp. A key to the 17 recog nized Caribbean species of Mycale is provided. Ectosomal skeletal patterns currently used as diagnostic characters for subgenera of Mycale may be inad equate for phylogenetic analysis, but reliable alternative congruent traits have not yet been identified to replace these. More than 150 species of Mycale Gray have strengthened the need for a better as have been described worldwide (Doumenc sessment of the genus' biodiversity, for a & Levi 1987), with representatives in most stable system of classification, and for bet marine habitats. They are common in both ter descriptions to differentiate between al polar and tropical seas, and have been re lopatric sibling species. ported from intertidal pools abyssal depths It has been convincingly argued that de (Hartman 1982). Contemporary mono scriptions of Mycale are often unreliable in graphs have added great numbers of new respect to noting size categories of micro species (Levi 1963, van Soest 1984, Berg scleres (Hentschel 1913, Doumenc & Levi quist & Fromont 1988, Hajdu & Desquey 1987). Accordingly, special attention was roux-Faundez 1994), indicating that many here paid to this important characteristic. more undescribed taxa are yet to be found. Details of rnicrosclere shape too were long Our study of the rich marine mangrove eco considered to be useful characters because system of Belize (Riitzler & Feller 1988, of their low adaptive value (Ridley & Den 1996; de Weerdt et al. 1991) is no exception dy .1887, Dendy 1921, Hajdu et al. 1994a, and has revealed two new species of Myc Hajdu & Desqueyroux-Faundez 1994). This ale. Recent findings of metabolites with view is adopted here and supported by our pharmacological potential from species of scanning electron micrography. Mycale (e.g., Capon & Macleod 1987; Per ry et al. 1988, 1990; Fusetani et al. 1989; Materials and Methods Corriero et al. 1989; Butler et al. 1991; Sponges were collected by the authors Northcote et al. 1991; Hori et al. 1993) and Kathleen P. Smith during several sur- 738 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON o, Twin Cays ..... ,'... ~~ :riJ"~': o ci iQj South Water Cay c!S9.....Blue Ground (;: Carrie Bow Cay ···n'!. ". Range ):r: (§:: : Ii CCurlew Bank !I! : . ....... (.: f ~ /ti.:...;.OJ t .:~ :::...... .... .. (:~~.:.: c........ ) L......J 1 krn r······......... Fig. I. Map of collecting area on the barrier-reef platform of Belize. Geographical coordinates for principal localities are 16°48.18'N, 88°04.93'W, Carrie Bow Cay; 16°49.95'N, 88°06.34'W, Twin Cays; and 16°48.55'N, 88°08.89'W, Blueground Range. veys of mangroves in the vicinity of Carrie scanning electron microscopy (SEM) were Bow Cay, Belize (16°48'N, 88°05'W) (Fig. prepared in one of the two following ways: I). Specimens are deposited in the sponge Samples in ethanol were washed of debris collection of the National Museum of Nat with several jets of ethanol, air dried in an ural History, Washington, subsamples were oven, and mounted on SEM stubs by ap donated to the Museu Nacional, Universi plying a thin layer of Entellan (Merck); dade Federal do Rio de Janeiro, Brazil. samples from ethanol were washed in warm Schizotypes of the two new species were water (ca. I min), immersed in a saturated also deposited in The Natural History Mu solution of soda, and heated to 50-70°C (30 seum, London, Queensland Museum, Bris min). Then they were again washed in bane, and Zoologisch Museum Amsterdam. warm water (1 min) and etched in 20% Most anatomical preparations were made HN03 at room temperature. After a few according to Riitzler (1978) and Hajdu minutes under observation, samples were (1994). Additionally, skeletons for study by rinsed with warm-water jets (1 min) and a VOLUME III, NUMBER 4 739 few drops of hydrogen peroxide (H20 2) so ston, 1842 (misspelled as H. aegogropila) lution added to the last change of water. Af as a type-species of Aegogropila (Thiele ter 30 min at 50-70°C the samples were 1903) overlooked Gray's (1867) name, Ae rinsed in warm water (1 min), transferred gogropila varians. Because Ae. varians was to ethanol (96%, 30 min), air-dried in an probably a replacement name in order to oven, and mounted on SEM stubs. avoid tautology, although not clearly stated Abbreviations used are: BMNH (The so by Gray (1867), we choose the figured Natural History Museum, London), INV specimen of H. aegagropila in Johnston POR (Invemar-Porifera Collection, Univer (1842, BMNH 1847.9.7.39) as lectotype of sidad Nacional de Colombia, Santa Marta), Ae. varians. In this way, both species be MCZ (Museum of Comparative Zoology, come objective synonyms. This point was Harvard University, Cambridge), MNHN made clear by L. B. Holthuis (in lit.). (Museum National d'Histoire Naturelle, The plasticity reported here for the pres Paris), MNRJ (Museu Nacional, Universi ence of ectosomal reticulation in Mycale dade Federal do Rio de Janeiro), MSNG (Ae.?) carmigropila new species is taken as (Museo Civico di Storia Naturale di Gen indication that some representatives of Car ova), MUT (Museo della UniversiHi di To mia (see below) are more closely related to rino), NNM (National Natuurhistorisch Mu Aegogropila than to other species of Car seum, Leiden), QM (Queensland Museum, mia, suggesting paraphyly of the former Brisbane), UFRJPOR (Universidade Fed subgenus and polyphyly of the latter. Ad eral do Rio de Janeiro, Porifera collection), ditionally, some M. (Mycale) species may USNM (National Museum of Natural His have their confused tangential ectosomal tory, Washington), and ZMA POR (Zoolo skeleton developed so thinly as to make gisch Museum Amsterdam, Porifera collec spicules strewn at random appear reticulat tion). ed to the casual observer (cf. M. [M.] thielei Hajdu & Desqueyroux-Faundez 1994). Systematic Descriptions Species like this have been assigned to Ae gogropila in the past, for instance M. (M.) Order Poecilosclerida Topsent, 1928 fiagelliformis (Bergquist & Fromont, 1988) Suborder Mycalina Hajdu et al., 1994 which has pore-grooves and a confused ec Family Mycalidae Lundbeck, 1905 tosomal skeleton. It becomes apparent that Genus Mycale Gray, 1867; sensu Hajdu & the monophyletic status of Aegogropila Desqueyroux-Faundez 1994 may not hold up in a thorough revision of Diagnosis.-Mycalidae with skeleton of all species. Such a revision is not yet pos subtylostyles and palmate anisochelas; ad sible, given that collections are dispersed, ditional microscleres may include sigmas, many new taxa are still being discovered, toxas, micracanthoxeas, raphides, unguifer and, more importantly, no other congruent ate anisochelas, and palmate isochelas. characters are yet apparent to replace the existing system. Subgenus Aegogropila Gray, 1867 Mycale (Aegogropila) arndti van Soest, Diagnosis.-Mycale with tangential ec 1984 tosomal skeleton of reticulate spicule tracts Figs. 2, 3, 17a; Table 1 (often easily peeled off), without serrated sigmas. Type species: Aegogropila varians Esperia macilenta.-Carter 1871 :276, pI. Gray, 1867 (=M. [Ae.] contarenii [von 17, fig. 8; not Hymeniacidon macilenta Martens, 1824, as Spongia contarenii.]). Bowerbank, 1866 (=Mycale (Carmia) Remarks.-Bergquist & Fromont (1988) macilenta). in quoting Halichondria aegagropila John- Mycale macilenta.-Arndt 1927:143, in 740 PROCEEDI GS OF THE BIOLOGICAL SOCIETY OF WASHI GTO Fig. 2. Mycale amdri, skeleton: a, Peeled-off surface showing ectosol11al reticulation; scale equals 500 1'-111; b, Detail of surface reticulation showing sigmas-I disposed around the bundles of megascleres, and disposition of rosettes (R) in the corners of the meshes; scale equals 100 I'-m; c, d, Fusiform subtylostyles and all four categories of anisochelas (I-IV); scale equals 1001'-111. (a-c, US M 39329: d, holotype ZMA POR 3675.) part (not the figured specimen); not Hy Description.-Macroscopic appearance: meniacidon maciLenta Bowerbank, 1866 Grayish blue to gray crust or cushion with (=MycaLe (Carmia) maci/enta). oscules often raised like small chimmneys; M. (Aegogropi/a) amdti van Soest, 1984: non-slimy when rubbed. Preserved material 19, pI. II, fig. 1. Zea 1987:140. is composed of seven main fragments, the MateriaL studied.-ZMA POR 3675, ho largest is 55 mm long, 30 mm wide, and up to 4 mm thick. The color in alcohol is drab lotype, Cura~ao; ZMA POR 3842, paratype, Florida. USNM 43032, Belize, Twin Cays, or pink, varying from light pink to almost Batfish Point, < 1 m, colI. K. Riitzler, 27 violet. The fragments have firm consistency Apr 1989; USNM 39329 (MNRJ 630), Be and microhispid surface texture. The ecto lize, Blue Ground Range, 1 m, colI. E. Haj some has sand grains embedded in the sur du & K. Smith, 12 Aug 1993; USNM face and is easily peeled off. No oscules are 47871, Blueground Range, on red-man apparent. grove stilt root, < 1 m, colI. K. Smith, Nov Skeleton: The ectosomal skeleton is a 1996. tangential reticulation of 120 f.Lm thick sub Diagnosis.-Bluish, crustose Aegogro tylostyle bundles (Fig.