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. 2a), with no appar­ pi/a with sigmas (three categories) and tox­ ent cementing spongin. Meshes are mostly as; four categories of anisochelas, with cat­ triangular and up to 400 X 200 f.Lm in di­ egories II and IV having exceptionally long ameter. Microscleres occur in abundance frontal alae (alae of head and foot nearly and may be playing the structural role of fused), covering the entire shaft in frontal holding the reticulation intact. Sigmas I (as view. many as 10 per linear mm) and rosettes VOLUME III, NUMBER 4 741

(135 fJ-m, 6 per mm) of anisochelas I can tial if a morphological-phylogenetic classi­ be seen surrounding the megasclere bundles fication of MycaLe is to be achieved. In (Fig. 2b). These and other microscleres, oc­ many taxonomic descriptions, one or the cur in great numbers inside the meshes. other microsclere category is often over­ The choanosome area appears light looked or lost to lumping decisions, as in brown in transmitted light, with dark-brown the present species where anisochelas-cat­ blotches of fibrous spongin. The spiculation egory III was not recognized by previous here is not very dense and no clear pattern authors (van Soest 1984, Zea 1987). Con­ is apparent which may be due to the slightly sequently, anisochelas II of M. arndti macerated condition of the specimen. Mi­ should be described as arcuate because the croscleres are scattered throughout this lateral alae of the head are partly separated area. from the shaft (Hajdu et al. 1994). Spicules (Figs. 2c, d, 3; Table 1): Sub­ The stout sigmas I are comparable to tylostyles fusiform, gradually tapering to a those of EspereLLa simonis as figured by point, generally slightly bent in upper third Ridley & Dendy (1887) and of M. quadri­ and thickest at mid length, with long neck partita Boury-Esnault (Hajdu & Desquey­ and eliptical head. Anisochelas I stout, with roux-Faundez 1994), and may be homolo­ head length 50% of total length, foot in side gous to the diancistras of Hamacantha Gray view at about 110° angle to shaft; frontal (Hajdu 1994). This similarity is based On alae of foot may bear denticulation on top the straight inner face and the abrupt taper­ (Fig. 3e). Anisochelas II similar to aniso­ ing of the hooks in this type of sigma and chelas I but even stouter, with head length is enhanced by a faintly marked notch at 70% of total length, shaft not visible in mid-length of some of these spicules (Fig. frontal view, lateral alae of head arcuate. 3h). Toxas show a remarkable range of size Anisochelas ill slender, with head length but size-frequency analysis of 100 toxas in 70% of total length, shaft slightly bent at the Belizean specimen failed to reveal dis­ the end of the lateral alae of the head. An­ tinct categories. isochelas IV very slender, with both frontal alae prolonged into thin, digitiform pro­ MycaLe (AegogropiLa?) carmigropiLa, cesses that sometimes cross each other. Sig­ new species mas I stout, abruptly bent into very sharp Figs. 4, 17b; Table 2 hooks, with almost straight inner faces. Sig­ mas II slender, with gradual curve and Type materiaL.-Holotype, USNM sharp hooks. Sigmas ill similar to sigmas 34560, Twin Cays, Sponge Haven, on roots II but smaller. Toxas very gently bent (more of Rhizophora, 1 m, colI. E. Hajdu & K. pronounced in short forms). Smith, 17 Aug 1993. Paratypes: USNM EcoLogy.-Associated with mangrove 42997, Belize, Twin Cays, Batfish Point, on and shallow seagrass bottoms in bays and red-mangrove stilt root and peat bank, < 1 lagoons. m, colI. K. Riitzler and K. Smith, 5 May Distribution.-Florida, St. Vincent (Vir­ 1986; USNM 43048, Belize, Twin Cays, gin Islands), Curac;:ao, Colombia, Belize. Hidden Creek, on red-mangrove stilt root, Remarks.-Only one of this species <1 m, colI. K. Riitzler, 8 May 1987; USNM specimen was found, suggesting that it may 43047, Belize, Twin Cays, Hidden Creek, be uncommon in the survey area. Details of encrusting a cluster of algae, genus HaLi­ spicule morphology and presence of four meda, <1 m, colI. K. Smith, 8 May 1987; types of anisochelas agree well with the ho­ USNM 47870, Belize, Twin Cays, Sponge lotype (Figs. 2d, 3b-d). Haven, peat bank with algae, genus HaLi­ It should be emphasized that proper as­ meda, 1 m, colI. K. Smith, 5 luI 1990; sessment of microsclere categories is essen- USNM 34561, Twin Cays, Sponge Haven, 742 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Mycale amdti, spicules: a, Face view of anisochela-I; scale equals 10 IJ.m; b, Face view of anisochela­ II showing robustness and extremely long frontal ala of head, almost fusing with foot; note deeply concave top portion of frontal ala of the foot (arrow); scale equals 10 IJ.m; c, Oblique frontal view of anisochela-III; scale equal 5 IJ.m; d, Profile view of anisochela-IV showing thin digitiform prolongations from both frontal alae (arrows) almost fusing; scale equals 5 IJ.m. e, Detail of foot of anisochela-I in Fig. 3a showing tridentate (t) top VOLUME Ill, NUMBER 4 743 on roots of Rhizophora, 1 m, colI. E. Hajdu other two specimens USNM 34561, 38768 & K. Smith, 17 Aug 1993; USNM 34587, lack the specialized tangential ectosomal Twin Cays, Sponge Haven, on roots of Rhi­ skeleton, having just a few scattered me­ zophora, 1 m, coll. E. Hajdu & K. Smith, gascleres, rosettes, and abundant sigmas 17 Aug 1993; USNM 38768, Twin Cays, near the surface. Sponge Haven, on Halimeda, 1 m, colI. E. The choanosomal skeleton is relatively Hajdu & K. Smith, 17 Aug 1993. Schizo­ low in spicule density, which accounts for types from holotype, BMNH 1994.3.1.1, the extreme softness and fragility. Subty­ QM G313152, MNRJ 631, ZMA paR lostyles occur scattered in great numbers 10708; fragments of paratypes, MNRJ 632, and also form longitudinal bundles (:530 633,634. fLm wide), frequently meandriform, that end Diagnosis.-Intensely blue to greenish, in fan-like brushes (:5500 fLm wide) in the encrusting MycaLe (Aegogropila?) with subectosome. These brushes support the three categories of anisochelas and one type tangential ectosomal reticulation, when of small «70 J..Lm) sigmas. present, or the rosettes alone; generally, fi­ Description.-Macroscopic appearance: bers protrude slightly from the surface of 2 Specimens were encrusting (:520 cm ) on the sponge. roots of Rhizophora, or occurred intermin­ Spicules (Fig. 3, Table 2): Subtylostyles gled with HaLimeda. In the first case they slender, slightly fusiform, straight or slight­ were cobalt-blue, while in the second they ly bent in central part, with eliptical head appeared very light green. Other specimens well marked, shaft gradually tapering to a observed alive were blue to violet, turning sharp point. Anisochelas I with straight tan when dying. Subectosoma1 canals con­ shaft, head 40% of total length; frontal alae verging on oscules in a star-like pattern are forming narrow angle (30°) with shaft (pro­ visible in live specimens. The consistency is very soft, slimy, and fragile, the texture file view), frontal alae of feet lower than rather smooth. lateral alae, general aspect narrow in both Skeleton: The ectosomal skeleton is a frontal and profile views. Anisochelas II polymorphic feature in this species. Two shaft markedly bent at head 50% of total specimens (USNM 34560, 34587) have a length, lateral alae of head longer than fron­ tangential reticulation of subtylostyles, sin­ tal alae. Anisochelas III very slender, with gle or in 50 fLm thick bundles, with large shaft gradually curved in profile view, head amounts of cementing spongin. Meshes are 40-50% of total length, frontal alae of foot mostly triangular (250 X 150 fLm). Up to distally extended by angled spurs. Sigmas 12 pores (40 X 20 to 80 X 50 fLm in di­ with markedly bent hooks, gradually sharp­ ameter) of the aquiferous system can be ening ends. seen within a single mesh. A few rosettes EtymoLogy.-The name carmigropiLa is of anisochelas I (:5 110 J..Lm in diameter), derived from the juxtaposition of Carmia and sigmas I occur scattered among the and AegogropiLa. These are the two sub­ meshes, frequently around the bundles. The genera with which specimens of this spe-

portion of frontal ala; scale equals 10 IJ.m; f, Oblique view from the back of an anisochela-II showing arcuate­ like pattern exhibited by the lateral alae of its head, which are elongated (arrow), rather than cut-off from the shaft; scale equals 5 IJ.m; g, Profile view of c1eistochela-like anisochela-IV showing thin digitiform prolongations from both frontal alae intercrossing each other (arrows); scale equals 5 IJ.m; h, S-shaped, diancistra-like sigma­ I showing almost straight inner-face of hooks (h), and rare, notch-like constriction (n) at tortion point; scale equals 20 IJ.m; i, Sigma-l1; scale equals 5 IJ.m; j, Sigma-III; scale equals 5 IJ.m; k, Toxas of stable geometry, but variable length; scale equals 20 IJ.m. (a, e-k, USNM 39329; b-d, holotype ZMA POR 3675.) 744 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Mycale carmigropila, spicules: a, Terminations of subtylostyles; scale equal LO fl.m; b, Face view of ani ochela-I howing narrow aspect; scale equals 10 fl.m; c, Profile view of anisochela-I; scale equals 10 fl.m. d, ProfiLe view of ani ochela-II showing narrow aspect, and shaft markedly bent at middle portion; scale equals 5 fl.m; e, Profile-view of anisochela-nI, note short, and thin digitiform spur on top of frontal ala of foot (arrow); scale equals 5 fl.m; f, Sigma; scale equals 20 fl.m. (a, d-f, Aegogropila-like ho)otype, USNM 34560; b, c, Carmia­ like paratype, US M 34561.) VOLUME Ill, NUMBER 4 745

N cies show affinities in ectosomal skeletal II patterns. -5- r­ Ecology.-On Rhizophora mangrove stilt I l/"l roots and Halimeda algae bordering a large mangrove channel with strong water flow. ~ r- ., Distribution.-Belize. "t:: II o Remarks.-Of the four specimens stud­ :; .A Q. ~ ~ ~ l/"l ied, two have Aegogropila-like ectosomal Vi J, I oc skeletons and two lack it, thus resembling '" Carmia (Arenochalina species are also without ectosomal skeleton but instead have =: ~ 11 oi a stout choanosomal reticulation). Both ~ 1 forms are indistinguishable in the field and C'I have identical spiculation. The assignment -a c -00 to Aegogropila is tentative but justified by -I V> 1- C the view that the ectosomal reticulation is '""lr)" .,co ~I more likely to be lost in a species of Ae­ E _C'I ...!.X gogropila than acquired spontaneously in 0\ one of Carmia. Mycale (Aegogropila?) carmigropila is '"I close to Mycale (Carmia) magnirhaphidi­ N- -I lera (see below) except for the possession of one more category of anisochelas, slight differences in the shape of anisochelas and lack of raphides and of a second category of sigmas. Both species may have blue or bluish color in life.

0) "t:: o Mycale (Aegogropila) citrina, new species Q. -5- ~ Figs. 5, 6, 17c; Table 3 C'" C'I o C'I c Mycale americana van Soest, 1984:22 (in part).-Zea 1987: 138. Type material.-Holotype, USNM .... 38942, Twin Cays, Cuda Cut, 1 m, colI. E. .E Hajdu & K. Smith, 13 Aug 1993. Paratypes, E l/"l ::l. l/"l USNM 43031, Belize, Twin Cays, Cuda "'00 3 II Cut near Batfish Point, encrusting a cluster ""'"....; " J1 of algae, genus Halimeda, < 1 m, colI. K. c "''9 ., '7'" E S;x Riitzler, 27 Apr 1989; USNM 38956, ~ C'I :::l 38958, 38963, 38967, Twin Cays, Cuda V> .,co Cut, 1 m, colI. E. Hajdu & K. Smith, 11 E 0) Aug 1993; USNM 47872, Belize, Twin "'5 u Cays, Cuda Cut, peat bank, < 1 m, coil. K. "5. Riitzler, 10 May 1997; USNM 41443 Be­ '1 lize, Twin Cays, Cuda Cut, peat bank, < 1 m, coll. K. Riitzler, 10 May 1997. Schizo­ types from holotype, BMNH 1994.3.1.2, QM G313153, MNRJ 635, and ZMA POR 746 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

10709; fragments of paratypes, MNRJ 636­ M e- 00 639. ~~~ Diagnosis.-Lemon-yellow crustose Ae­ l()oQ~ "i'''i'''i' gogropila with three categories of aniso­ N N­ lrl lrl lrl chelas, one type of large sigmas (>60 /-Lm), and copious mucus production when han­ dled. Description.-Macroscopic appearance: 5' M~~ N Thick incrustations (5 rom; 30 cm2) with II r!-r!-rJ, protruding oscules encircled by a thin mem­ ::~~ III brane. Extremely soft and fragile, and re­ ONM leasing copious amounts of mucus upon c handling. Live-color is lemon yellow to

C light orange, turning pale yellow in alcohol. '"co Q) Numerous, pronounced subectosomal chan­ E nels may cover most of the sponge surface and converge toward the oscules. Most specimens contained embryos during Au­ gust. Skeleton (Fig. 5): The ectosomal skele­ ton peels off easily .and is made of a re­ markably pure (no scattered megascleres) tangential reticulation of bundled (~85 /-Lm thick) subtylostyles (Fig. 5a), mostly form­ ing triangular meshes (660 X 400 /-Lm), with no apparent cementing spongin. Ro­ settes of anisochelas-I (130 /-Lm in diameter) occur in places (Fig. 5b, c). Sigmas are very abundant, scattered among the meshes (up to 35 per mesh), and along the strands (30 per rom; Fig. 5d); they seem to play an im­ portant structural role in the skeleton. The choanosome is crowded with dark­ brown granules (about 40 J.Lm in diameter), ... - presumably of mangrove peat, which ham­ ..e -~ e­ per observation of the skeletal architecture. E N ::l. The underlying pattern seems to be densely e, spicular, with crisscrossing megasclere bun­ c'" dles (150 /-Lm thick) and abundant micro­ Q) scleres. Ee ::l Spicules (Fig. 6, Table 3): Subtylostyles '"co Q) straight, slender and slightly fusiform, with E poorly marked eliptic head. Anisochelas I with straight shaft, head 50% of total length. Anisochelas-II with shaft bent at head, head 50-60% of total length; frontal alae· of fOGI: with shGrt, digitif-eFIR··pr-eoesses­ on top (Fig. 6c). Anisochelas-III very slen­ der, with shaft gradually cur.ved (profile VOLUME 111, NUMBER 4 747

Fig. 5. Mycale citrina, skeleton: a, Peeled-off surface showing ectosomal reticulation; scale equals 500 j.1m; b, Detail of surface reticulation showing sigmas disposed abundantly around the bundles of megascleres, and rosette of anisochelas-l; scale equals 200 j.1m; c, Detail of rosette of anisochelas-I shown on below left corner of Fig. 5b; scale equals 50 j.1m; d, Detail of bundle of megascleres showing abundance of sigmas around it; scale equals 50 j.1m. (a-d, paratype USNM 38963.) view) head 40% of total length, foot with 1987). Reexamination of Zea's material re­ very small alae, the frontal one ending on vealed the occurrence of an intermediate­ top in a thin digitiform process (Fig. 6d). size category of anisochelas, overlooked by Sigmas slender, markedly bent on hooks. the author. This feature and the larger di­ Etymology.-The name citrina is derived mensions of spicules in the Colombian ma­ from citrus, for the lemon-yellow color of terial make us confident in assigning the the live sponge. Colombian specimens to M. citrina. Ecology.-Specimens were found under Only one specimen quoted by van Soest the roofs of mangrove overhangs, or, less (1984) in the type-series of Mycale ameri­ commonly, on peat banks where they were cana was not collected in a mangrove protected from direct sunlight by neighbor­ (ZMA POR 3889, on Halimeda). Unfortu­ ing bushy algae (e.g., species of Halimeda, nately the live color of the specimen was Caulerpa, Jania). not registered but comparison with the ho­ Distribution.-Belize. lotype (Table 3) revealed three categories of Remarks.-This species is close to Myc­ anisochelas, instead of two as originally ale americana van Soest (Fig. 7), but dif­ quoted, thicker megascleres, and slightly fers by the possession of a third category larger sigmas. We suggest to assign this of anisochelas, larger megascleres and sig­ specimen to M. citrina. mas, and lemon-yellow color. M. americana was originally described as red (van Soest Subgenus Arenochalina Lendenfeld, 1887 1984), but orange-yellow and olive-yellow Diagnosis.-Mycale without ectosomal specimens have also been reported (Zea skeleton, with choanosomal skeleton of 748 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Mycale citrina, spicules: a, Terminations of subtylostyles; scale equals 10 !LID; b, Face view of anisochela-I; scale equals 5 !Lm; c, Profile view of anisochela-II, note short and thin digitiform spur on top of frontal ala of foot (arrow); scale equals 5 !LID; d, Profile view of anisochela-III, note thin digitiform prolongation on top of poorly developed frontal ala of foot (arrow); scale equals 5 !Lm; e, Slender sigma; scale equals 10 !Lm. (a-e, holotype USNM 38942.) stout fibers forming a coarse rectangular re­ ing established, as suggested by similar oc­ ticulation, and with simple spicule comple­ currences here and there in Mycale assigned ment. Species of this subgenus are known to other subgenera. to exude large amounts of mucus upon han­ dling. Type species: Arenochalina mirabilis von Lendenfeld, 1887 (=M. mirabilis, sen­ Mycale (Arenochalina) laxissima su Wiedenmayer 1989). (Duchassaing & Michelotti, 1864) Remarks.-The status of Arenochalina is Figs. 8, 9, 17d; Tables 4, 5 uncertain. The diagnosis is in essence that provided by van Soest (1984) for Acama­ Acamas laxissima Duchassaing & Michel­ sina de Laubenfels, 1936a. Wiedenmayer's otti, 1864:95, pI. XXII, fig. 3. (1989) addition with respect to the frequent Esperella nuda Ridley & Dendy, 1886:339; coring of fibers by foreign debris is not up­ 1887:70, pI. XV, figs. 5, 11, 14; pI. XVI, held here as this seems to be noteworthy fig. 1. for Australian species only. Shared traits Hircinia cartilaginea.-Hyatt 1877:549 among populations from both sides of the (not Spongia cartilaginea Esper, 1798). Atlantic, the Indo-west Pacific and southern Hircinia cartilaginea (Esper) var. horrida Australia, seem to be the choanosomal, Hyatt, 1877:549; pI. 17, fig. 29. coarse, rectangular reticulation of spiculo­ Hircinia purpurea Whitfield, 1901:49, pI. 4 fibers, the low diversity of categories in the (not H. purpurea Hyatt, 1877:550). spicule complement, and the production of Mycale angulosa.-De Laubenfels 1936a: abundant mucus. Nonetheless, the derived 116, fig. 2, pI. 15, 1.-? Levi 1959:129; condition of these characters is far from be- fig. 19; pI. 6 fig. 5; Lopez & Green 1984: o-< S -~ -z ~ Table 3.-Spicule measurements (in lim) for Mycale citrina and Mycale americana (ranges of length, or length X width, with means in italics; n = 20, unless ~ stated). .I:>-

Material Subtylostyles Anisochelas I Anisochelas II Anisochelas III Sigmas

Mycale citrina Holotype USNM 38942 (MNRJ 635) 339-382.4-456 X 8-8.6-9 32-45.5-48 19-21.6-24 13-/3./-14 70-74.9-79 Paratypes USNM 38963 (MNRJ 636) 376-4/8.4-461 X 8-9.1-10 41-45.5-54 20--21.4-24 12-12.6-14 69-77.7-88 USNM 38956 (MNRJ 637) 350-427.4--477 X 7-8.6-11 44-49.4-52 19-23.2-26 12-12.9-14 75-80.4-88 USNM 38958 (MNRJ 638) 408-422.9-456 X 8-9.2-11 43-49.7-54 21-23.0--25 12-/3.2-15 72-78.3-86 USNM 38967 (MNRJ 639) 366-399./-424 X 7-8./-10 45-49.2-54 18-21.0-23 12-/2.7-14 76-78.7-85 M. americana (Zea 1987) 309-341.1-371 X 3-7.1-9.5 35-40.4-47 not reported 10-//.6-13 60--66.9-70 INV-POR 0241 (remeasured) 318-328.9-366 X 5-6.9-9 33-38.4-41 21 (n = 1) 10-13 63-68.4-74 X 3-4 INV-POR 0276 (remeasured) 276-311.1-350 X 2-4.5-7 33-36.0-40 16-19 (n = 5) 11-13 58--64.9-72 X 2-4 M. americana (van Soest 1984) 190-265.6-338 X 5-7.2-10 30-36.9-40 10-/3.9-22 not reported 23-47.6-63 Holotype (ZMA POR 4074, remeasured) 196-242.9-260 X 5--6.8-8 34-36.9-40 not found 12-13.3-14 32-47.0-52 Paratype (ZMA POR 3889, remeasured) 276-309.5-329 X 7-9.0-11 34-37.6-42 18-20.6-22 11-12.4-14 54--61.4--67

-...) .I:>­ '0 750 PROC~EDINGSOF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Spicules of Mycale americana, for comparison with Mycale citrina (Fig. 6): a, Terminations of subtylostyles: scale equals 10 J.Lm; b, Profile view of anisochela-I; scale equals 5 J.Lm; c, Profile view of aniso­ chela-III, note absence of digitiform process on top of well developed frontal ala of foot (compare with Fig. 6c); scale equals 5 J.Lm; d, Stout sigma; scale equals 10 J.Lm. (a-d, holotype ZMA POR 4074.)

79 (not Pandaros angulosa Duchassaing Mycale jamaicensis Pulitzer-Finali, 1986: & Michelotti, 1864:89; pI. IX fig. 4). 125. Mycale sp.-Reiswig 1973. Mycale whitfieldi Pulitzer-Finali, 1986: 127. Thorecta horridus.-Wiedenmayer 1977: Mycale hyatti Pulitzer-Finali, 1986:129. 70, pI. 8, figs. 2-4; pI. 9, fig. 1. Acamasina laxissima.-Wiedenmayer Material studied.-MUT POR 34, holo­ 1977:146-147, 255. type, St. Thomas, Virgin Islands. MSNG, Mycale laxissima.-van Soest 1981:12.­ no register number; Jamaica, Port Royal, on van Soest et ai. 1983:200.-Pulitzer-Fin­ submerged ruins, 5-10 m, 27 Mar 1964 (M. ali 1986:1l9.-Riitzler 1990:455.-Va­ laxissima sensu Pulitzer-Finali, 1986; num­ celet 1990:25. ber PRo 40); MSNG 47695, Jamaica, Port ?Mycale impeifecta.-Winterman-Kilian & Royal cays, 10-25 m, 22 Mar 1964 (M. mu­ Kilian 1984: 133. cijiuens Pulitzer-Finali, 1986; holotype); Mycale (Acamasina) laxissima.-Van Soest MSNG 47697, Jamaica, Duncans, fore-reef 1984:29; pI. III, 1; fig. 9.-Zea 1987:143, slope, 35 m, 30 Mar 1964 (M. jamaicensis 255. Pulitzer-Finali, 1986; holotype); ZMA POR Mycale mucifluens Pulitzer-Finali, 1986: 5192, Jamaica, Runaway Bay, 33.5 m, 14 121. Aug 1969, colI. and det. H.M. Reiswig Mycale nuda.-Mello-Leitao et ai. 1961: (Mycale sp.). MCZ 7008/cat. 440, Florida, 12.-Hechtel 1976:254.-van Soest Key West; Hircinia cartilaginea sensu Hy­ 1984:31.-Hajdu & Boury-Esnault 1991: att, 1877 (M. hyatti Pulitzer-Finali, 1986; 504. holotype); MCZ 70711cat. 441, Florida, VOLUME Ill, NUMBER 4 751

Fig. 8. Mycale laxissima, skeleton: a, Detail of skeleton showing fused fibers (f), and meshes of widely variable dimensions (m); scale equals 500 J.Lm; b, Detail of skeleton showing primary(l) and secondary (2) interconnecting fibers; scale equals 500 J.Lm; c, Stout megascleres with blunt apex, and anisochelas, young megasclere is seen below center; scale equals 50 J.Lm; d, Anisochelas of variable geometry; scale equals 20 J.Lm.

Key West; H. cartilaginea sensu Hyatt, diameter, and thin-walled (0.5-2 cm). The 1877 (M. hyatti Pulitzer-Finali, 1986; para­ color is dark wine red. A large pseudoscule type); MCZ 7073/cat. 222, Florida, Cape (6 cm diameter) on top of the tubes is en­ Florida; H. cartilaginea var. horrida Hyatt, circled by a transparent membrane. The 1877; M. hyatti Pulitzer-Finali, 1986; para­ smaller specimens (about 6 X 6 X 6 cm) type. UFRJPOR 3578, Brazil, Angra dos are globular and already bear an apical Reis, Rio de Janairo State. USNM 41273, pseudoscule. The sponge is tough but elas­ Belize, Twin Cays, Cuda Cut near Batfish tic, compressible, spiny to the touch, and Point, <1 m, colI. K. Riitzler, 4 Jun 1983; releases copious amounts of sticky mucus USNM 39281 (several lots), Belize, Twin when squeezed. The surface is coarsely Cays, Cuda Cut, 1 m, colI. E. Hajdu and K. conulose from projecting spiculo-fibers. Smith, 11 Aug 1993. Skeleton (Fig. 8a, b): The surface mem­ Diagnosis.-Dark red, spiny, tubular Ar­ brane can be peeled off in places and con­ enochalina with stout spiculo-fibers envel­ tains a few scattered spicules and paucispic­ oped by great amounts of spongin and ular strands of subtylostyles. The choano­ forming large rectangular meshes. Choan­ somal skeleton is made up of a stout, rect­ osome cavernous, without noticeable fleshy angular reticulation of spiculofibers. parts, with only one category of anisochelas Meshes formed are generally within the and large (>50 /-Lm) sigmas. range of 100-150 /-Lm up to 3-4 mm (Fig. Description.-Macroscopic appearance: 6a). One can distinguish between primary Specimens are tube shaped, some in clus­ and secondary fibers (50-300 /-Lm thick; ters up to six, up to 50 cm tall, 15 cm in Fig. 8b), the latter showing clearer spongin 752 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 9. Mycale laxissima, spicules: a, Rare larger, and common smaller anisochelas; scale equals 10 ILm; b, Common anisochela of intermediary size; scale equals 10 ILm; c, Common anisochela of smaller size; scale equals 10 ILm; d, Stout sigmas, and anisochelas of intermediary size; scale equals 20 ILm. in transmitted light. The fibers end as sub­ mas occur in abundance, strewn at random ectosomal, paucispicular tufts. Anisochelas inside the skeleton meshes. are particularly abundant in a subectosomal Spicules (Figs. 8c, d, 9; Table 4): Sub­ layer of spongin where they are spread out tylostyles are straight, variably thick, without order. Rosettes of anisochelas are abruptly tapering to a blunt apex, with seen on the basal plate of spongin in a spec­ ovoid head. Anisochelas are variable in di­ imen from Angra dos Reis, Rio de Janeiro mensions and geometry, generally appear State, Brazil; UFRJPOR 3578. Sigmas are narrow in profile and face views, have a very common along the spiculofibers of the curved shaft and a head 40-50% of total choanosome. Isolated megascleres and sig- length. Sigmas are generally stout and

Table 4.-Spicule measurements (in ILm) for Mycale laxissima (fragments of different specimens) from Belize (ranges of lengths, or length x width, with means in italics; n = 10).

Material SubtylostyJes Anisochelas Sigmas

USNM 39281 (MNRJ 640) 270-283.6-302 x 5--6.5-8 20-23.9-31 83-94.5-106 X 4--6 MNRJ 641 249-272.4-292 X 5--6./-10 22-23.6~29 83-93.3-112 X 2-3.2-6 MNRJ 642 249-277.2-297 X 6--6.5-9 18-24.5-30 79-96.1-108 X 3-5 MNRJ 643 260-279.8-297 X 3-7.5-9 21-27.2-32 80-96.7-112 X 4-6 MNRJ 644 249-28/.7-307 X 5--6.9-9 22-27.6~33 87-96.3-105 X 4-4.8-7 MNRJ 645 244-262.9-286 X 4--6.6-8 18-24.8-32 90-100.9-114 X 6 MNRJ 646 265-285.7-302 X 5--6.3-8 21-25.8-32 83-94.3-110 X 4-6 USNM 41273 265-285.1-302 X 5-5.9-7 19-25.6-34 87-97.0-112 X 4-6 o-< l' c:::: ~ tr1 - ~ Table 5.-Spicule measurements (in /-Lm) for Mycale laxissima from Belize compared to other Caribbean locations (ranges of length, or length X width, with means tr1 in italics; n = 20, unless stated). ~ .j>.

Material Subtylostyles Anisochelas Sigmas Raphides

Belize (population, n = 80) 244-278.6-307 X 3-6.5-10 18-25.4-34 79-96.1-114 X 3-7 not found Holotype (MUT 34IUSNM 31001, remeasured) 224-248.5-272 X 3-4 14-18.6-21 not found not found Holotype (cf. Pulitzer-Finali 1986) 230-260 X 2.5-3.5 16-20 50-71 (rare) not reported van Soest 1984 209-223.2-240 X 2-3.5 16-20.0-23 70-75.0-81 not reported Pulitzer-Finali 1986 220-250 X 3.5-4.5 20-23 69-80 X 3.5 not reported remeasured 227-249.9-269 X 4-4.5-6 21-22.4-23 73-79.5-84 X 3-4 not found Zea 1987 (Providencia) 219-276.0-304 X 4-5.7-6 16-20.1-23 75-83.7-93 X 2-2.8-4 not reported Zea 1987 (Santa Marta) 223-254.1-271 X 2-4.3-6 16-19.7-21 72-78.3-85 X 2-2.8-4 not reported Zea 1987 (Cartagena) 242-294.5-323 X 3-8.1-11 26-30.2-33 109-118.3-128 X 5-6.2-9 not reported Zea 1987 (San Bernardo) 213-279.3-313 X 3-9.0-11 25-29.9-36 109-//8.7-133 X 6-6.7-8 not reported Ridley & Dendy 1887 (as Esperella nuda) 245 X 16 25 120 not reported remeasured 235-259.8-283 X 5-9.0-13 22-26.1-30 76-/00.9-112 X 4-6 not found Pulitzer-Finali 1986 (as M. mucifiuens) 250-290 X 3.5-4.5 19-29 77-93 X 2-3.5 not reported remeasured 258-278.2-294 X 3-4.5-6 21-25.1-29 84-88.8-95 X 2-3.4-4 not found Pulitzer-Finali 1986 (as M. jamaicensis) 250-280 X 3-3.5 16-23 65-90 X 2-2.5 100 (rare) remeasured 249-266.4-283 X 3-4.4-6 15-19.5-25 76-80.0-87 X 2-2.8-3 not found Pulitzer-Finali 1986 well, I (as M. whitfieldi) 214-270 X 2.5 18.5-22.5 71-76 not reported Pulitzer-Finali 1986 (as M. hyatti) 230-300 X 3-11.5 23-26 75-105 X 7.5 not reported remeasured 224-276.8-322 X 4-6.8-11 23-24.8-29 78-88.8-106 X 4-4.4-6 not found

-..J Ul W 754 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON abruptly bent into hooks with sharp end­ Red Sea and Indian Ocean, respectively, be­ ings. cause our examinations suggest that there is Ecology.-Moderately common in shad­ greater intra-than interspecific variation in ed marine mangrove environments with this taxonomic complex, for instance, of an­ strong tidal water flow, on protected patch isochelas in a single specimen. Clarification reefs below 5 m, and in fore-reef habitats of this problem will ultimately depend on below 12 m (compare also the detailed study of populations from many localities analysis of reef populations by Reiswig worldwide and we caution from new de­ (1973) in Jamaica (as Mycale sp.). The scriptions based on poorly preserved or un­ heavy spongin fibers are known to be bored representative material (e.g., related new and occupied by filamentous algae (Rutzler taxa in Pulitzer-Finali 1986). 1990) and experimental work by one of us The growth form of Mycale laxissima is (KR) has shown that the coring of siliceous generally tube or funnel shaped. In Belize, spicules contribute photosynthetically use­ even in shallow water, neither encrusting ful light levels inside the dark sponges, nor cushion-shaped specimens were seen comparable to glass-fiber light guides in op­ and early globular stages develop directly tical applications (similar observations for into long, thin-walled tubes. This is in con­ other siliceous sponges were reported by trast to observations in Brazil (between la­ Gaino & Sara 1994, and Cattaneo-Vietti et tidudes 8°-24°S; EH) where encrusting al. 1996). forms prevail among shallow-water popu­ Distribution.-Florida, Bahamas, Cuba, lations of this species. The consistency of Jamaica, Puerto Rico, Virgin Islands, Neth­ specimens varies, suggesting polymorphism erlands Antilles, Colombia, Belize, Mexico, in patterns of skeleton reticulation. Like­ Brazil, West Africa. wise, abundance of microscleres varies Remarks.-This species was subject of among specimens but no attempt was made several misinterpretations in the literature. to quantify differences within and between Esperella nuda Ridley & Dendy, 1886 from individuals because these sponges were of­ Brazil (holotype BMNH 1887.5.2.171) ten large and only fragments have been col­ turns out to be a junior synonym and the lected to allow cut specimens to regrow to originally reported (Ridley & Dendy 1887) protect their apparently small population exceptional thickness of its megascleres (16 size. f-Lm) was found to be only in the range of 5-13 f-Lm. Specimens described by Pulitzer­ Subgenus Carmia Gray, 1867 Finali (1986; as M. hyatti) and Zea (1987) possess comparable megascleres (4-11 f-Lm Diagnosis.-Mycale without ectosomal and 2-11 f-Lm thick; Table 5), an isolated skeleton, with loose choanosomal skeleton characteristic that does not justify separat­ composed mainly of plumose, isolated spi­ ing a species. The species was also con­ culofibers which are often paucispicular. fused with Mycale angulosa (de Laubenfels Type species: Hymeniacidon macilenta 1936a, Levi 1959),- as discussed by van Bowerbank, 1866 (=M. [C.] macilenta sen­ Soest (1984). Another erroneous identifi­ su Topsent 1924). cation may have been the report of M. im­ Remarks.-The absence of an ectosomal peifecta Baer from Colombia (Winterman­ skeleton is likely to be a homoplastic oc­ Kilian & Kilian 1984). The suggestion of currence in Mycale because it is found in possible amphi-Atlantic distribution of M. all species of Carmia and Arenochalina, laxissima (van Soest 1984) raises the ques­ and possibly also in the subgenus Mycale, tion of apomorphies of other distant spe­ (for instance, M. (M.?) lapidiformis (Ridley cies, such as M. setosa (Keller 1889) and & Dendy), Hajdu & Desqueyroux-Faundez M. euplectellioides (Row 1911) from the 1994). As such, it is a weak character on VOLUME 111, NUMBER 4 755

Fig. 10. Mycale magnirhaphidijera, spicules: a, Set of spicules showing raphides-I (arrows mark the extrem­ ities); scale equals 50 iLm. b, Terminations of tylostyles, note blunt apex, and well-marked oval head; scale equals 10 iLm. (a, b, USNM 39278.) which by itself to base a subgeneric diag­ call. K. Riitzler, 26 Apr 1989 (purplish wine nosis. red; sigmas moderately common and clear­ ly in 2 size classes); USNM 43040, Belize, Mycale (Carmia) magnirhaphidifera Twin Cays, Hidden Creek, < 1 m, call. K. van Soest, 1984 Riitzler, 1 May 1989 (gray, dirty yellow in­ Figs. 10, 11, 17e; Table 6 side and where dying, enveloping sabellid tubes; without sigmas); USNM 43033, Be­ M. (C.) magnirhaphidifera van Soest, 1984: lize, Twin Cays, Hidden Creek, < 1 m, call. 27; pI II, 7. K. Riitzler, 1 May 1989 (gray; sigmas abun­ M. cecilia.-Wells & Wells, in Wells et al. dant); USNM 43036, Belize, Twin Cays, 1960:212 (not M. cecilia de Laubenfels, Gator Creek, < 1 m, call. K. Riitzler, 3 May 1936b:447). 1989 (grayish ochre; sigmas very abun­ Material studied.-ZMA POR 4885, ho­ dant); USNM 43037, Belize, Twin Cays, lotype, Cura<;ao. USNM 23633, North Car­ Gator Creek, < 1 m, call. K. Riitzler, 3 May olina, Hatteras Harbor, call. and del. H. W. 1989 (grayish ochre, massive; sigmas very & M. J. Wells (as M. cecilia), 21 Nov 1959. abundant; with embryos); USNM 39278, USNM 42878, Belize, Twin Cays, Lair Belize, Twin Cays, Hidden Creek, 1 m, call. Channel, on red-mangrove stilt root, < 1 m, E. Hajdu & K. Smith, 9 Aug 1993 (bluish call. K. Riitzler, 10 May 1985 (dark royal­ purple, sigmas common); USNM 39279, blue, sigmas rare); USNM 42979, Belize, Belize, Twin Cays, Cuda Cut, 1 m, call. E. Twin Cays, Main Channel, on red-man­ Hajdu & K. Smith, 11 Aug 1993 (bluish grove stilt root, < 1 m, call. K. Riitzler and purple, sigmas common; with larvae); K. Smith, 26 Apr 1986 (purplish, with sa­ USNM 39280, Belize, Twin Cays, Turtle bellid worm tubes; many sigmas); USNM Pond, 1 m, call. E. Hajdu, 17 Aug 1993 42949, Belize, Twin Cays, Main Channel, (bluish purple, sigmas common); USNM on red-mangrove stilt root, < 1 m, call. K. 41442, Belize, Twin Cays, Cuda Cut, on Riitzler and K. Smith, 26 Apr 1986 (dark red-mangrove stilt root, <1 m, call. K. Riit­ wine red, tan below surface; no sigmas); zler, 2 May 1994 (grayish blue, very thin USNM 47874, Belize, Twin Cays, Grouper crust; sigmas very rare); USNM 47865, Be­ Gardens entrance, on a strand of rope tied lize, Twin Cays, Hidden Creek, < 1 m, call. to a plastic pipe protruding from the chan­ K. Riitzler, 29 Mar 1997 (blue, sigmas nel bottom, 1 m, call. K. Riitzler, 1 Feb abundant). 1986 (ochre, sigmas rare); USNM 43029, Diagnosis.-Bluish-purple to brown­ Belize, Twin Cays, Sponge Haven, 1 m, gray, crustose Carmia with two types of an- 756 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 11. Mycale magnirhaphidijera, spicules: a, Profile view of anisochela-I, note basal portion of lateral and frontal alae of the head diverging from each other (arrows), the lateral ones projecting behind the shaft; scale equals 5 fLm; b, Face view of narrow anisochela-I, and part of anisochela-II on top; scale equals 5 fLm; c, Profile view of anisochela-II showing short, thin digitiform spur on top of frontal ala of its foot (arrow); scale equals 5 fLm; d, Sigmas-I and -II; scale equals 10 fLm. e, Raphides-I; scale equals 5 fLm; f, Raphides-II; scale equals 5 fLm. (a, b, d, e, USNM 39278; c, f, holotype ZMA POR 4885.) -< ~ m ...... z ~ t;O m

.j>. Table 6.-Spicule measurements (in fLm) for Mycale magnirhaphidijera (ranges of length, or length X width, with means in italics; n = 20, unless stated). '"

Material Subtylostyles Anisochelas I Anisochelas n Sigmas I Sigmas II Raphides I Raphides II Cura~ao Holotype (ZMA POR 4885) 236-254.2-270 X 1.5~2.2-3 19-27.6-33 10-12.4-17 not found not found 260-291.3-310 8-9.4-11 North Carolina Wells & Wells 1960 (USNM 23633, as M. cecilia) 250-270 X 4-6 30-36 14-16 50-58 not reported not reported not reported remeasured 196-231.0-258 X 4-6 30-30.9-35 13-15.6-18 46-49.1-55 not found 168-204 (n = 7) 11-20 (n = 4) Belize USNM 39278 (MNRJ 647) 196-258.2-286 X 4.4 31-35 (n = 4) 11-13 32-43.6-50 13-21 (n = 3) 90-143.4-241 13-17.5-22 MNRJ 648 232-266.6-294 X 4.4-6.0 21-24.4-35 12-13.4-15 26-31 (n = 2) not found 280-304.9-322 15-17.0-19 MNRJ 649 232-278.0-308 X 3.9-5.0 29-36 (n = 5) 9-11.3-13 45-47.1-52 not found 112-189.8-249 13-20.9-28 MNRJ 650 258-283.1-308 X 3.9-5.5 29-31.6-34 10-11.2-13 37-43.7-47 19-23.0-26 120-153.7-260 14-19.3-37 MNRJ 651 232-269.4-283 X 4.4-5.3 31-32.7-36 11-14.1-16 40-43.1-48 19(n=l) 95-154.3-252 11-14.1-16 MNRJ 652 274-303.0-322 X 4.4-6.6 33-37 (n = 5) 11-11.9-18 42-45.0-47 20 (n = 1) 109-192.6-255 15-18.5-21 USNM 39280 (MNRJ 653) 255-279.2-300 X 4.2-5.4 31-32.8-36 11-12.0-13 40-43.2-46 not found 129-214.8-286 13-16.4-19 MNRJ 654 238-275.0-294 X 4.2-6.1 31-31.8-34 12-12.7-13 37-44.9-48 22 (n = 1) 101-213.1~280 14-19.3-23

--.l U. --.l 758 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON isochelas, with or without sigmas, and with II delicate with sharp points. Raphides I two size classes of raphides, the larger ones slender, straight, sharply pointed. Raphides reaching more than 200 ILm in length. II stout, straight, sharply pointed. Description.-Macroscopic appearance: Ecology.-A common species in the sur­ Encrusting, up to (5 mm thick) very soft vey area where it prefers shaded substrates. and fragile. Subectosomal channels of the Many specimens were found on the bare aquiferous system converging to oscula (::53 tips of new Rhizophora mangle prop roots mm diameter) which are raised and sur­ suggesting capacity to quickly colonize rounded by transparent collars. Color in life newly available substrate (R-strategist). The is highly variable (see material section common association with polychaete above), burgundy to bluish-purple and worms was also observed by Wells et al. grayish blue in light-exposed areas, cream (1960, as M. cecilia) and van Soest (1984). to yellowish in shaded zones; drab in dying Distribution.-North Carolina, Cura9ao, tissue parts. Most specimens collected (May Belize. to August) contained larvae (150-300 ILm Remarks.-In contrast to the holotype in diameter) in the choanosome; many were from Cura9ao, most Belizean specimens associated with clusters of serpulid poly­ contain abundant sigmas (see material sec­ chaete tubes. tion above). On the other hand, color, Skeleton: The ectosome, lacking a spe­ shape, and morphology of all spicules other cial skeleton, consists of a superficial mem­ than sigmas agree well, including the sec­ brane bearing pores (30-50 ILm diameter) ond smaller category of raphides that was and is supported by terminal brushes of found in the holotype (Fig. 11f) but was not choanosomal spiculofibers. A few sigmas originally reported (van Soest 1984). Nev­ and single anisochelas are seen between the ertheless, the abundance of sigmas in some pores, rosettes of anisochelas I (ca. 70 ILm specimens is striking (see material section diameter) occur in the subectosomal zone. above) and may justify separating the sig­ The choanosome has a low spicule den­ ma-bearing population as a different species sity, with subtylostyles organized in pauci­ if more supporting data become available. spicular (3 to 10) ascending tracts of 15-30 The material from North Carolina iden­ ILm in diameter. The fibers branch seldomly tified by Wells & Wells, in Wells et al. and generally run parallel to each other (1960) as Mycale cecilia has previously without anastomosing. Close to the surface been referred to both M. microsigmatosa they diverge to form brushes. Sigmas are (Hechtel 1965) and M. americana (van very common throughout the sponge. Soest 1984). We have analyzed the speci­ Raphides too are dispersed, not organized men (USNM 23633) and found it conspe­ into trichodragmas. cific with M. (C.) magnirhaphidifera, also Spicules (Figs. 10, 11; Table 6): Subty­ bearing sigmas (Table 6). Wells & Wells, in lostyles straight, slender, blunt, with char­ Wells et ai. (1960) described their material acteristic irregularly-oval head. Anisochelas as pale yellowish-green or yellowish-tan, I, narrow in face and side views, head about similar to the color of M. (C.) magnirha­ 50% of total length, base of frontal ala of phidifera from shaded habitats. the head curved forward, base of lateral alae of the head curved backward and Mycale (Carmia) microsigmatosa Arndt, slightly protruding over the shaft (in side 1927 view). Anisochelas II slender, head about Figs. 12, 13, 17f; Table 7 50% of total length, frontal ala of foot ex­ tended on top in a short digitiform process. Mycale fistulata var. microsigmatosa Arndt, Sigmas I with almost straight inner face, 1927:144, pI. I, 9. abruptly bent into very sharp hooks. Sigmas Mycale microsigmatosa.-Burton 1956: VOLUME III, NUMBER 4 759

Fig. 12. Mycale microsigmatosa, spicules: a, Terminations of subtylostyles showing heads, and gradually sharpening apex; scale equals 10 f.l.m; b, Terminations of subtylostyles showing head and gradually sharpening apex; scale equals 10 f.l.m. Ca, USNM 33580; b, holotype ZMA POR 1593.)

129.-Hechte1 1965:47.-van Soest Description.-Macroscopic appearance: 1981:12.-Winterman-Ki1ian & Kilian Specimens are encrusting or cushion­ 1984: 132.-Pulitzer-Finali 1986:124.­ shaped (5 cm thick), very soft and fragile. Riitzler 1986:120. Subectosomal channels of the aquiferous Mycale (Carmia) microsigmatosa.-van system converge to scattered oscula «5 Soest 1984:24; pUI, fig. 6.-Zea 1987: mm) with transparent, membranous collar. 142. Not sensu Green & Gomez 1986: Live color is highly variable but predomi­ 284 (=M. cecilia de Laubenfels). nantly reddish orange. Even grayish-green­ Carmia microsigmatosa.-Hajdu & Boury­ ish specimens have recognizable bright or­ Esnault 1991:510. ange specks which aid in field recognition Material studied.-ZMA POR 1593, ho­ of the species. Specimens with larvae were lotype, Cura~ao. USNM 22207, ho1otype of found during August. M. cecilia, Panama City, Panama. USNM Skeleton: A thin surface membrane with 33580, Belize, Twin Cays, Turtle Pond en­ scattered megascleres and a few paucispi­ trance, on root of Rhizophora, colI. I. cular (3 to 6 spicules wide) bundles can be Goodbody, 24 Feb 1985; USNM 34625, peeled off and terminal spicule brushes of Belize, Twin Cays, colI. I. Goodbody, Feb the ascending choanosomal fibers reach and 1986; USNM 41258, Belize, Twin Cays, penetrate the ectosome. Main Channel south of dock, on mangrove The choanosome is supported by mean­ roots, colI. K. Riitzler, 12 Iun 1983; USNM dering tracts of subtylostyles (50 /-Lm thick) 39302 (several lots), Belize, Twin Cays, which may be more or less replaced by for­ Cuda Cut, colI. E. Hajdu & K. Smith, 13 eign structures, such as algae and sabellid Aug 1993; USNM 39326 (severa11ots), Be­ polychaete tubes. These tracts split up and lize, Twin Cays, Sponge Haven, colI. E. anastomose near the surface where they fan Hajdu & K. Smith, 9 Aug 1993; USNM out in brushes. Scattered megascleres are 47873, Belize, B1ueground Range, <1 m, common throughout and sigmas are seen colI. K. Riitz1er, 6 May 1994. USNM here and there in large concentrations. 42951, Belize, Twin Cays, Batfish Point, Spicules (Figs. 12, 13; Table 7). Subty­ <1 m, colI. K. Riitz1er, 19 Apr 1996. lostyles slender, mostly straight, with faint­ Diagnosis.-Orange-dotted to reddish, ly marked oval head, apex sharpening grad­ crustose Mycale (Carmia), with two cate­ ually or abruptly. Anisochelas I, head about gories of small «30 /-Lm) anisochelas and 60% of total length, shaft gradually curved, small «50 /-Lm) sigmas. basal portion of frontal and lateral alae 760 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 13. Mycale microsigmatosa, spicules: a, Profile view of anisochela-I; scale equals 5 fLm; b, Profile view of anisochela-II; scale equals 5 fLm; c, Profile view of anisochela-II showing thin digitiform process on top of frontal ala of foot (arrow); scale equals 5 fLm; d, Slender sigma; scale equals 10 fLm; e, Profile view of anisochela­ I; scale equals 5 fLm; f, Profile view of anisochela-II showing thin digitiform process on top of frontal ala of foot (arrow); scale equals 5 fLm; g, Stout sigma; scale equals 10 fLm. (a, b, d, USNM 33580; c, holotype ZMA POR 1593; e-g, holotype of M. cecilia, USNM 22207.) o<: t ~ -tT'l -z ~ tl:I ~ Table 7.-Spicule measurements (in ,.,.m) for Myca/e microsigmalosa (ranges of length, or length X width, with means in italics; n = 20, unless stated). ...

Material Subtylostyles Anisochelas I Anisochelas II Sigmas

Holotype (ZMA POR 1593, Arndt 1927) 250-294 X 4.8 8-19 not given 32-38 remeasured 232-260.8-283 X 4-6 16-18.0-20 13-/3.3-14 30-32.4-37 USNM 33580 224-241.9-263 X 4-6 18-19.3-21 13-14.1-15 30-35.0--42 USNM 34625 213-237.4-252 X 6-7 18-19.0-20 12-14.0-15 25-33.8-36 USNM 41258 216-24/.6-260 X 3--4 18-18.6-20 13 (n = 3) 30-38.3--45 USNM 39302 (MNRJ 655) 235-253.8-274 X 3-5 18-/9.5-20 13-14 (n = 3) 29-33.8-37 MNRJ 656 224-236.9-249 X 2-4.1-5 17-18.9-21 12-13.5-15 34-,-35.0-37 MNRJ 657 216-239.7-277 X 3-4.1-6 17-19.6-21 14-/4.3-16 30-33.2-35 MNRJ 658 244-269.6-288 X 4-4.7-7 19-20.3-24 12-/3.8-15 33-36.9--41 MNRJ 659 235-247.0-263 X 4-6 18-18.7-20 13-/3.8-15 32-34.8-39 MNRJ 660 246-267.7-288 X 4-6 18-20.6-23 14-/4.8-15 31-36.6--42 MNRJ 661 238-252.0-273 X 3-5 18-18.8-20 12-13.4-15 32-35.7-39 MNRJ 662 227-240.5-258 X 3-4.2-6 17-18.2-20 13-13.4-15 36-37.8-39 MNRJ 663 232-251.7-267 X 3-4.2-6 17-18.6-20 11-13 (11 = 2) 31-34.4-39 USNM 39326 (MNRJ 664) 243-266./-288 X 4-6 18-19.5-21 13-/3.8-15 32-37.0--43 MNRJ 665 230-240.2-259 X 4-5.0-7 18-20.2-21 12-14 (n = 6) 31-35.9-39 MNRJ 666 224-255.4-274 X 3-4.5-6 18-20.0-23 13-14.6-16 32-34.7-36 MNRJ 667 221-249.2-269 X 3--4 18-/9.0-20 13-14.1-15 32-35.5-39

-.I -0- 762 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

slightly divergent. Anisochelas II, head of two categories of anisochelas in M. mi­ about 50% of total length, shaft gradually crosigmatosa rather supports the former curved, with basal portion of frontal and view, agreement with M. cecilia (holotype, lateral alae of head slightly divergent; in Fig. Be-g). Furthermore, de Laubenfels some of these spicules the top of the frontal (1936b) describes M. cecilia as "basically ala of the foot tapers to a thin digitiform green, abundantly provided with bright red process (Fig. 13f). Sigmas slender, both specks", similar to our own observations ends markedly bent into sharp hooks. on M. microsigmatosa. However, we prefer Ecology.-Common on mangrove prop to keep M. cecilia as a valid name for east­ roots, including new growth tips, and all ern Pacific records of the sponge, basing other light-exposed substrates in shallow la­ this decision on geographic separation and goon and harbor habitats (R-strategist). The small differences in spicule dimensions. species is also reported as tolerant of oil and Therefore we propose to assign the record domestic wastes (Muricy 1989). Sabellid of M. (C.) microsigmatosa from the Pacific polychaetes are commonly associated w.ith coast of Mexico (Green & Gomez 1986) to and overgrown by this sponge. M. cecilia, this sponge has relatively thick Distribution.-Bermuda, Florida, Baha­ subtylostyles (5-9 /-Lm) and anisochelas mas (?), Cuba (?), Jamaica, Puerto Rico, with a large range of length (12-21 /-Lm), Venezuela, Netherlands Antilles, Colombia, possibly representing two categories. Van Belize; also from Brazil, West Africa (7), Soest (1984) also suggested possible syn­ A~ores (?), and Eastern Mediterranean (7). onymy of Mycale microsigmatosa with M. Remarks.-Agreement with the holotype senegalensis Levi, 1952, M. sanguinea (Figs. 12b, 13c) is excellent, including the Tsurnamal, 1969, and Desmacella melior­ presence of two categories of anisochelas ata Wiedenmayer, 1977. To this list we may which has not before been reported for the add Desmacella janiae Verril, 1873, Biem­ species. Because anisochelas are rare in this na microstyla de Laubenfels, 1950a, and species (van Soest 1984, Hajdu & Boury­ Prosuberites scarlatum Alcolado, 1984, but Esnault 1991), it is not surprising that oc­ formal decision must await re-examination currence of a second category was previ­ of all types. ously overlooked. Mycale microsigmatosa is very common Subgenus Mycale Gray, 1867 in Belizean mangroves, displaying mainly reddish-orange color but also red and trans­ Diagnosis.-Mycale with ectosomal lucent gray. Similar color variability was skeleton of confused tangential megascleres reported by Hechtel (1965, Jamaica), van over perpendicular, dendritic choanosomal Soest (1984; Cura~ao, Bonaire, Margarita, spicule tracts that end in subectosomal and Florida), and Zea (1987, Colombia). brushes. Type species: Hymeniacidon lin­ Hajdu & Boury-Esnault (1991) found Bra­ gua Bowerbank, 1866 (Hajdu & Desquey­ zilian specimens (collected between lati­ roux-Faundez 1994). tudes 13° and 24°S) to be exclusively red Remarks.-A subgenus Anomomycale but subsequent observations (EH) found was erected by Topsent (1924) for Desma­ pale-yellow specimens from Angra dos cidon titubans Schmidt (1870). This Reis (Rio de Janeiro state). sponge, however, has most characteristics Bergquist (1965) and Hechtel (1965), of the subgenus Mycale, including the su­ suggested the synonymy of Mycale micro­ bectosomal brushes, large megascleres (one sigmatosa with M. cecilia de Laubenfels. size class >700 /-Lm), and possibly tangen­ Another opinion by van Soest (1984) fa­ tial subtylostyles in the ectosome (Lund­ vored a close relationship between M. ce­ beck 1905). Only the distinctive shape of cilia and M. americana. The present finding its anisochelas separates it. This single fea- VOLUME Ill, NUMBER 4 763

Fig. 14. Mycale laevis, skeleton: a, Transverse section showing terminally divergent spicule brushes (s) of ascending choanosomal tracts supporting the tangential ectosomal skeleton (e); scale equals 200 /Lm; b, Detail of surface showing tangentially disposed megascleres, and abundance of rnicroscleres; scale equals 100 /Lm; c, Detail of subectosomal area showing typical patches of trichodragmas, and also of isolated raphides; scale equals 50 /Lm; d, Fusiform, bent subtylostyle, note abundance of trichodragmas; scale equals 100 /Lm. ture is insufficient to justify subgeneric sep­ j.Lm) megascleres and two size classes of aration and it seems appropriate to keep M. anisochelas; with basal, spur-like projec­ titubans in the subgenus Mycale. tions on the smaller anisochelas; also pres­ ent, two classes of sigmas and raphides in Mycale (Mycale) laevis (Carter, 1882) trichodragmas. Figs. 14, 15, 17g; Table 8 Description.-Macroscopic appearance: Thick cushions (up to 5 cm) or clusters of Synonymy, see Hajdu & Desqueyroux­ 20 or more branches (1-10 cm diameter, up Faundez (1994); in addition: to 50 cm tall) rising from a massive base Mycale jUsca.-Mello-Leitao et al. 1961: and often with apical oscula (up to 4 cm 12; Hechtel 1976:254; Hajdu & Boury diameter). Oscula have fibrous-looking col­ Esnault 1991:504. lars made up of tracts of megascleres. Con­ Mycale (Mycale) laevis.-Lehnert 1993:54. sistency is rough but compressible and the Material studied.-USNM 41267, Be­ texture rough. Color in life is a bright or­ lize, Twin Cays, Sponge Haven, < 1 m, on ange-yellow, but also a few whitish speci­ mangrove roots, coll K. Rtitzler, 13 June mens were seen. Specimens with embryos 1983; USNM 39328 (3 lots), Belize, Blue were seen in August. Ground Range, 1 m, colI. E. Hajdu & K. Skeleton (Fig. 14a-c): The ectosome is Smith, 12 Aug 1993. reinforced by a thin layer of tangentially Diagnosis.-Orange to yellow, encrust­ strewn subtylostyles interspersed with ing or massive Mycale with large (>500 abundant microscleres. A very dense, con- 764 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. IS. Mycale laevis, spicules: a, Face view of anisochela-I; scale equals 20 fLm; b, Profile view of anisochela-I, note slight s-shape of shaft; scale equals 20 fLm; c, Detail of foot of anisochela-I shown in Fig. 15b, with an undulated basal portion of the falx; scale equals 5 fLm; d, Face view of rare, robust anisochela-III showing thin digitiform prolongation on top of frontal ala of foot (arrow up), and basal spur (arrow down); scale equals 5 fLID; e, Profile view of typical anisochela-III showing basal spur (arrow), note reduced foot without alae; scale equals 5 fLm; f, Sigmas-I and -II; scale equals 10 fLm; g, Trichodragmas-I and -II; scale equals 20 fLm; h, Rare trichodragmas-II; scale equals 5 fLm. VOLUME 111, NUMBER 4 765

fused subectosomal reticulation of multi­ = spicular subtylostyle tracts (up to 300 !-Lm thick) is oriented mainly parallel to the sur­ face. Spaces between tracts are filled by large amounts of scattered subtylostyles, anisochelas I, and patches of trichodragmas I (by far the most common microscleres, Fig. l4c). Brushes of subtylostyles support the ectosomal network (Fig. 14a), and pierce the surface slightly. Anisochelas I form a few rosettes (160 !-Lm) but more of­ ten occur scattered or surrounding the bun­ dles of megascleres.

II The choanosome shows high spicule -$ density and includes criss-crossing subty­ N lostyle tracts with loose subtylostyles and ~ 00 patches of numerous trichodragmas I in be­ - tween. Digitiform processes of the larger specimens are supported by longitudinal spicule tracts and a secondary, transverse plumo-reticulate skeleton. Spicules (Figs. 14d, 15; Table 8): Sub­ tylostyles fusiform, mostly bent with faintly marked, oval head and abruptly sharpened, or strongyloid apex. Anisochelas type I, 00 N with shaft slightly s-shaped, head ca. 40% -~ ~ of total length, basal portion of falx on foot oci oci-. 'I I 00 1'- undulated (Fig. 15c). Anisochelas type III - - slender, head 70% of total length, foot often ~ ~ reduced (lacking alae) and with basal spur. 00 00 J, ....,I Sigmas I stout, with sharply bent hooks ~ ~ Sigmas II similar to I but more slender. Tri­ 00 1'- 1'- chodragmas I; trichodragmas II. Ecology.-This is a well known reefal species (Goreau & Hartman 1966) and is widely distributed in the shallow parts (16­ 25 m) of the fore-reed of Carrie Bow Cay (see Rtitzler & Macintyre 1982 for the bion­ omy of the reef). In the mangroves, Mycale laevis is restricted to areas with good ex­ posure to the open, near-reef lagoon (it is common at Blue Ground Range and Cat Cay Lagoon, but very rare at Twin Cays). There, however, the sponges attached to red mangrove roots can grow to record size. Specimens laying unattached on the bottom (presumably fallen from the roots) and in­ tergrown with turtle grass survive well and appear healthy. 766 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Distribution.-Cuba, Jamaica, Domini­ USNM 38967, Belize, Twin Cays, Cuda can Republic, Puerto Rico, Venezuela, Co­ Cut, 1 m, colI. E. Hajdu & K. Smith 13 Aug lombia, Belize, Mexico; also Brazil. 1993. Remarks.-A few details should be add­ Spicules.-Sigmas (Fig. 16), slender, 0­ ed to former descriptions of the species. or s-shaped, shaft subcylindrical (flattened Notably, we recognize two categories of in cross section), with ca. 10 spines each sigmas (smaller one can be rare) rather than along the outer bends of the hooks; spines a large size range of one, and two catego­ single or double and directed toward spic­ ries of trichodragmas in some of the spec­ ule center. imens (again, the smaller being rare see Remarks.-These distinctive spicules are Hajdu & Desqueyroux-Faundez 1994, tab. the only material found, thus making spe­ 1, and references therein). No taxonomic cies diagnosis impossible. The sigmas came importance is given to these characters in abundance from a very small specimen here. or fragment (used up in making the prepa­ Esperia massa Schmidt, 1862 sensu ration) inadvertently collected as part of a Schmidt 1870:58 and Mycale massa specimen of M. citrina. (Schmidt, 1862) sensu Winterman-Kilian Comparison with the only Paresperella 1984:133, are likely to be synonyms of species described from the western Atlantic, Mycale laevis because the latter species was Mycale (P.) spinosigma (Boury-Esnault originally described from the Mediterra­ 1973; MNHN D NBE 968), indicates that nean Sea and a West-Indian distribution is the latter is most likely contaminated with highly unlikely despite the similarities in Paresperella spicules. The anisochelas re­ skeleton morphology. ported by Boury-Esnault (1973) were not found in a preparation of the type specimen Subgenus Paresperella Dendy, 1905 and sigmas were very rare. The ZMA col­ lection has a very small macerated speci­ Diagnosis.-Mycale with an ectosomal men (ZMA 5389) from inside Curavao skeleton of reticulated tangential mega­ (Barbara Beach, 1-3 m) which, from the scleres and with serrated sigmas among the appearance of serrated sigmas may be con­ microscleres. Type species: Esperia serra­ specific with the Belizean material. The Cu­ tohamata Carter, 1880. ravao specimen has slender subtylostyles Remarks.-Van Soest (1984), based on (270-323 /Lm long), one category of ani­ the very peculiar characteristics of serrated sochelas (24-30 /Lm long), and the serrated sigmas, considers Paresperella a monophy­ sigmas (78-98 /Lm long). One toxa (34 /Lm letic group. Nevertheless, in the absence of long) was also seen but may be foreign. We another congruent diagnostic character, Pa­ refrain from making an identification until resperella may fall to Aegogropila (see more material is available. comments on Anomomycale, above, under subgenus Mycale). However, until further study of species in Paresperella and other Discussion closely related subgenera (Aegogropila, Some names associated with Caribbean Carmia), it is convenient to maintain this Mycale are still in need of a reappraisal. subgenus for the 20 or so species already Winterman-Kilian & Kilian (1984) listed described with serrated sigmas. M. cleistochela fiagellifer Vacelet & Vas­ seur, 1971 (a species from Madagascar) in Mycale (Paresperella) species indet. Colombia, with sigmas twice the size of the Fig. 16 original report. We have not examined the Material studied.-Spicules found con­ Colombian material but assume that it is taminating sample of Mycale citrina, conspecific with M. (Ae.) diversisigmata VOLUME Ill, NUMBER 4 767

Fig. 16. Mycale (Paresperella) sp., characteristic sigmas (other spicules belong to M. citrina): a, Serrated sigma; scale equals 20 t-Lm; b, Serrated sigma showing non-cylindrical shaft, note narrower middle portion when compared to wider hooks; scale equals 20 t-Lm; c, Detail showing spines on hooks of serrated sigmas, note heterogeneous distribution with the occurrence of twins (arrows); scale equals 5 t-Lm. van Soest, 1984 (from Cura¥ao), which Schmidt, 1862 (sensu Schmidt 1870) is contains flagelliform sigmas (150-200 considered dubious by Topsent (1924), and ILm), as well as semi-closed anisochelas. unrecognizable by van Soest (1984). Win­ Similarity of the latter species with M. terman-Kilian & Kilian (1984) report M. cleistochela jiagellijer was suspected by massa from Colombia. However, data in­ van Soest (1984) who must have been un­ dicate that these specimens are conspecific aware of the morphology of true cleisto­ with M. laevis which has indeed very sim­ cheles which was only recently revealed by ilar spiculation. scanning electron microscopy (Pulitzer-Fin­ As argued above, Desmacidon titubans ali 1996:fig. 15; for M. (Ae.) peculiaris). described by Schmidt (1870) is best iden­ Schmidt (1870) cited five species from tified as Mycale (M.) titubans, although the Florida under genera Esperia Nardo, 1833 conspecificity of populations from Florida and Desmacidon Bowerbank, 1864, that (Schmidt 1870), the northeastern-Atlantic were referred to Mycale by subsequent au­ (Lundbeck 1905, Topsent 1924) and the thors. The specimen originally named Es­ Antarctic (Boury-Esnault & Van Beveren peria renieroides Schmidt (1870) was re­ 1982) is certainly unlikely and needs to be ferred to Oxymycale by van Soest (1984) confirmed. Esperia diaphana Schmidt and to Mycale by Pulitzer-Finali (1986). (1870; material examined: dissociated spic­ Neither author had examined Schmidt's ules mount made by Schmidt, BMNH preparation (BMNH 1870.5.3.31) which re­ 1870.5.3.32) and Esperia immitis Schmidt veals an isotropic reticulation of small ox­ (1870; material examined: dissociated spic­ eas, part of a haplosclerid sponge, and a ules mount made by Schmidt, BMNH single rosette of anisochelas of the type that 1870.5.3.30, MCZ No. 42; label could be is indicative of M. immitis (Schmidt 1870). in Schmidt's handwriting but depth of 274 The specimen identified as Esperia massa m differs from that of 230 m in the original 768 PROCEEDINGS OF niE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 17. Underwater photographs of live Mycale specimens representing seven of the eight species found at Twin Cays: a, M. arndti, grayish-colored specimen, X1.0; b, M. carmigropila new species, on red-mangrove stilt root, XO.9; c, M. citrina new species, on mangrove peat, X1.3; d, M. laxissima, on red-mangrove stilt roots, XO.3; e, M. magnirhaphidifera, overgrowing serpulid polychaete tubes on red-mangrove stilt roots, xO.5; f, M. microsigmatosa, X1.0; g, M. laevis on red-mangrove stilt root, XO.3. VOLUME 111, NUMBER 4 769 account) are regarded as good species of M. (C.) microsigmatosa this paper Mycale (Mycale). Esperia diaphana has Arndt two categories of megascleres (721-827 M. (Grapelia) ungui­ Hajdu et al. 1995 f.Lm and 509-604 f.Lm), three categories of fera Hajdu et al. M. (Mycale) diaphana Schmidt 1970 (as anisochelas (148-170 f.Lm, 31-40 f.Lm, 19­ (Schmidt) Esperia); this pa­ 25 f.Lm), one category of sigmas (16-23 per (discussion) f.Lm), and one category of trichodragmas M. (M.) immitis Schmidt 1970 (as (41-70 f.Lm), and seems close to M. (M.) (Schmidt) Esperia); this pa­ grandis Gray, 1867 (Indo-west Pacific), and per (discussion) M. (M.) anisochela Levi, 1963 (south and M. (M.) laevis (Carter) this paper southwestern Africa). Abundant material M. (M.) titubans Schmidt, 1870 (as from the Gulf of Mexico, agreeing with M. (Schmidt) Desmacidon); this (M.) diaphana is in the USNM collection. paper (comment, M. (M.) immitis has two categories of me­ subgenus Myca1e) M. (Paresperella) spe­ this paper gascleres (996-1198 f.Lm and 466:-530 f.Lm), cies indet. three categories of anisochelas (90-105 f.Lm, 29-33 f.Lm, 20-25 f.Lm), one category Key to the species of Caribbean Mycale of sigmas (35-45 f.Lm), and two categories 1. Megascleres near 500 /Lm common .. 2 of trichodragmas (90-108 f.Lm, 18-33 f.Lm). Megascleres always <500 /Lm 5 Van Soest (1984) considers Little's 2. Two categories of megascleres 3 (1963) Mycale macilenta from the Gulf of One category of megascleres, trichod- Mexico to be a likely new species in the ragmas often abundant . subgenus Aegogropila. In the account be­ · Mycale (M.) laevis low we will treat it as Mycale (Ae.) species. 3. Three categories of anisochelas, aniso- The following key to 17 recognized spe­ chelas I >60 /Lm 4 cies of Mycale of the Caribbean should be One category of anisochelas <60 /Lm, used in conjunction with the relevant liter­ twisted M. (M.) titubans 4. Anisoche1as I ::; 150 /Lm common .... ature (most recent reliable or available ref­ · M. (M.) diaphana erence). Anisochelas I <110 /Lm, with shaft Mycale (Aegropila) van Soest 1984 markedly curved in profile view ..... americana van Soest ·...... M. (M.) immitis M. (Ae.) angulosa van Soest 1984 5. Serrated sigmas present . (Duch. & Mich.) · M. (Paresperella) species indet. M. (Ae.) arndti van van Soest 1984 Serrated sigmas absent 6 Soest 6. Three or more categories of anisochelas 7 M. (Ae.) carmigropila this paper One or two categories of anisochelas . .. 12 new species 7. Sigmas present ...... 8 M. (Ae.) citrina this paper Sigmas absent, two categories of toxas new species · M. (Aegropila) species indet. M. (Ae.) diversigmata van Soest 1984 8. Two or more categories of sigmas ... 9 van Soest One category of sigmas ...... 10 M. (Ae.) escarlatei Haj- Hajdu et al. 1995 9. Two categories of sigmas, anisochelas du et al. II very narrow ("duck's-bill") . M. (Ae.) species indet. Little 1963 (as M. · M. (Ae.) escarlatei macilenta) Two categories of fiagelliform sigmas, M. (Arenochalina) lax­ this paper three of normal ones . issima (Duch. & · M. (Ae.) diversisigmata Mich.) Three categories of sigmas, four cate­ M. (Carmia) magnirha­ van Soest 1984 gories of anisoche1as .... M. (Ae.) arndti phidifera van Soest 10. Anisochelas only palmate 11 770 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Anisochelas I and II unguiferate, ani- ed by fellowships and grants from CNPq sochelas III palmate with spur . and FAPESp, Brazil. His field work in Be­ · M. (Grapelia.) unguifera lize was made possible by a short-term vis­ 11. Sigmas "?70 flom common, live sponge itor grant from the Smithsonian Office of lemon-yellow M. (Ae.) citrina Fellowships and Grants and by a fellowship Sigmas <70 flom, live sponge blue ... from the Smithsonian's Caribbean Coral · M. (Ae.) carmigropila 12. Raphides common...... 13 Reef Ecosystems program (CCRE; partly Raphides absent or very uncommon supported by a grant from the EXXON (possibly contaminants) ...... 14 Corporation), National Museum of Natural 13. Two categories of raphides, raphides I History. This is CCRE contribution number > 100 flom, isochelas absent ...... 531. · M. (Carmia) magnirhaphidifera One category of raphides <50 flom, is­ Literature Cited ochelas present ...... M. (Ae.) angulosa 14. With tangential ectosomal reticulation Alcolado, P. 1984. Nuevas especies de esponjas en­ · M. (Ae.) americana contradas en Cuba.-Poeyana 271: 1-22. Without tangential ectosomal skeleton 15 Arndt, w. 1927. Kalk- und Kieselschwamme von Cu­ 15. Sigmas common, >50 flom, coarse ra~ao.-Bijdragen tot de Dierkunde 25: 133­ choanosomal spongin fibers, massive 158, pis. I-III. sponges .... M. (Arenochalina) laxissima Bergquist, P. R. 1965. The sponges of Micronesia, part Sigmas <40 flom, delicate choanosomal 1. The Palau Archipelago.-Pacific Science 19: 123-204. skeleton strands, encrusting sponges .. ---, & J. P. Fromont. 1988. The marine fauna of (C.) · M. microsigmatosa New Zealand: Porifera, Demospongiae. Part 4 (Poecilosclerida).-New Zealand Oceanograph­ Acknowledgments ic Institute Memoir 96: 1-197. Boury-Esnault, N. 1973. Campagnes de la Calypso au We are indebted to E R. Schram and R. large des cotes atlantiques de I'Amerique du Sud (1961-1962). I. 29. Spongiaires.-Resul­ W. M. van Soest (Amsterdam) for critical tats scientifiques des Campagnes de la Calypso review of early drafts of the manuscript. L. 10:263-295. B. Holthuis (NNM) clarified doubts con­ ---, & M. van Beveren. 1982. Les Demosponges cerning the type species of AegogropiLa. A. du plateau continental de Kerguelen-Heard.­ B. Johnston (MCZ), C. Levi (MNHN), V. Comite National Fran~ais des Recherches An­ tarctiques (CNFRA) 52: 1-175. Rainieri (MSNG), S. Stone and C. Valen­ ---',& M. T. Lopes. 1985. Les Demosponges lit­ tine (BMNH), and S. Zea (lNV) are torales de l'archipel des A~ores.-Annales de thanked for the loan of specimens. C. Val­ l'Institut Oceanographique, nouvelle serie entine is specially thanked for her efforts 61(2):149-225. on behalf of Johnston's Halichondria ae­ Bowerbank, J. S. 1864. A monograph of the British Spongiadae. Vol. I: 1-290, pis. I-XXXVII. Ray gagropiLa. K. Smith (National Museum of Society, London. Natural History, Washington, D.C.) assisted ---. 1866. A monograph of the British Spongia­ in field work in Belize and in laboratory dae. Vol. 2: 1-388. Ray Society, London. preparations; she is credited for finding a Burton, M. 1956. The sponges of West Africa.-At­ new species, Mycale (Aegogropila?) car­ lantide Report. Copenhagen 4: 111~147. Butler, M. S., R. J. Capon, R. Nadeson, & A. A. Bev­ migropila. We thank M. K. Ryan (National eridge. 1991. Aromatic Bisabolenes from an Museum of Natural History, Washington, Australian Marine Sponge, Arenochalina sp.­ D.C.) for preparing Fig. I and helping with Journal of Natural Products 54:619-623. other illustration chores, and C. C. Hansen Capon, R. J., & J. K. Macleod. 1987. Structural and (also from the Museum) for taking and stereochemical studies on marine norterpene cy­ clic peroxides; part 2.-Journal of Natural printing the photograph in Fig. 17d. E Products 50:225-229. Hiemstra (Amsterdam) is thanked for ad­ Carter, H. J. 1871. On two undescribed sponges and vice on SEM preparation. EH was support- two Esperiadae from the West Indies; also on VOLUME Ill, NUMBER 4 771

the nomenclature of the calcisponge Clathrina, with the description of some new genera.-Pro­ Gray.-Annals and Magazine of Natural His­ ceedings of the Zoological Society of London tory (4)8: 1-28. 1867:492-558, pis. XXVII-XXVII. ---. 1880. Report on specimens dredged up from Green, G., & P. G6mez, 1986. Estudio taxon6mico de the Gulf of Manaar and presented to the Liv­ las esponjas de la Bahia de Mazatlan, Sinaloa, erpool Free Museum by Capt. W. H. Cawne Mexico.-Anales del Instituto de Ciencias del Warren.-Annals and Magazine of Natural His­ Mar y Limnologfa, Universidad Nacional Au­ tory (5)6:35-61. t6noma de Mexico 13(3):273-300. ---. 1882. Some sponges from the West Indies and Hajdu, E. 1994. A phylogenetic interpretation of ha­ Acapulco in the Liverpool Free Museum, de­ macanthids (Demospongiae, Porifera) with a re­ scribed with general and classificatory re­ description of Hamacantha popana (de Lauben­ marks.-Annals and Magazine of Natural His­ fels, 1935).-Journal of Zoology 232:61-77. tory (5)9:266-301, 346-368, pis. XI-XII. ---, & N. Boury-Esnault. 1991. Marine Porifera Cattaneo-Vietti, R., G. Bavastrello, C. Cerrano, M. of Cabo Frio (Rio de Janeiro, Brazil). The fam­ Sara, U. Benatti, M. Giovine, & E. Gaino. 1996. ily Mycalidae Lundbeck, 1905, with the de­ Optical fibres in an Anarctic sponge.-Nature scription of a new species.-Revista brasileira 383:397-398. de Biologia 51(3):503-513. Corriero, G., A. Madaio, L. Mayol, V. Picadilli, &·D. ---, & R. Desqueyroux-Faundez. 1994. A synop­ Sica. 1989. Rotalin A and B, two novel diter­ sis of South American Mycale (Mycale) (Poe­ pene metabolites from the encrusting Mediter­ cilosclerida, Demospongiae), with the descrip­ ranean sponge Mycale rotalis.-Tetrahedron 45: tion of three new species and a preliminary cla­ 277-288. distic analysis of Mycalidae.-Revue suisse de Dendy, A. 1905. Report on the sponges collected by Zoologie 101(3):563-600. Prof. Herdman at Ceylon in 1902.-Report to ---, R. W. M. van Soest, & J. N. A. Hooper. 1994. the government of Ceylon on the pearl-oyster Proposal of a phylogenetic subordinal classifi­ fisheries of the Gulf of Manaar 3 (Supplement cation of poecilosclerid sponges. Pp. 123-139 18):57-246, pis. I-XVI (Royal Society, Lon­ in R. W. M. van Soest, Th. M. G. van Kempen don). & J. C. Braekman, eds., Sponges in time and ---. 1921. The tetraxonid sponge spicule: a study space. Proceedings of the Fourth International in evolution.-Acta Zoologica V (2):95-152. Porifera Congress. Balkema, Rotterdam, 515 Doumenc, D., & C. Levi. 1987. Anisochelae analysis and taxonomy of the genus Mycale Gray (De­ pp. mospongiae). Pp. 73-92 in J. Vacelet & N. Bou­ ---, S. Zea, M. Kielman, & S. Peixinho. 1995. ry-Esnault, eds., Taxonomy of Porifera from the Mycale escarlatei n.sp. and Mycale unguifera northeast Atlantic and Mediterranean Sea. n.sp. (Mycalidae, Poecilosclerida, Demospon­ NATO ASI Series, vol. G13. Springer-Verlag, giae) from the tropical western Atlantic.­ Berlin, 332 pp. Beaufortia 45: 1-16. Duchassaing de Fonbressin, P., & G. Michelotti. 1864. Hartman, W. T. 1982. Porifera. Pp. 640-666 in S. P. Spongiaires de la mer Caraibe. Natuurkundige Parker, ed., Synopsis and classification of living Verhandelingen van de hollandsche Weten­ organisms (I). MacGraw-Hill, New York, 1166 schappen te Haarlem (2)21 (III): 1-124, pis. 1­ pp. XXv. Hechtel, G. J. 1965. A systematic study of the De­ Esper, E. J. C. 1791-1830. Die Pflanzenthiere. Raspe, mospongiae of Port Royal, Jamaica.-Peabody Niirnberg. Museum of Natural History Bulletin 20:1-103. Fusetani, N., N. Yasumuro, S. Matsunaga, & K. Hash­ ---. 1976. Zoogeography of Brazilian marine De­ imoto. 1989. Mycalolides A-C, hybrid macro­ mospongiae. Pp. 237-260 in E W. Harrison & lides of ulapualides and halichondramide, from R. R. Cowden, eds. Aspects of sponge biology. a sponge of the genus Mycale.-Tetrahedron Acaderrlic Press, New York, 354 pp. Letters 30:2809-2812. Hentschel, E. 1911. Tetraxonida 2. Teil. Pp. 277-393 Gaino, E., & M. Sara. 1994. Siliceous spicules of Teth­ in W. Michaelsen, & R. Hartmeyer, eds., Die ya seychellensis (Porifera) support the growth Fauna Siidwest-Australiens 3 (10). Gustav Fi­ of a green alga: a possible light conducting sys­ scher, Jena, 467 pp. tem.-Marine Ecology Progress Series 108: ---. 1913. Uber einen Fall von Orthogenese bei 147-151. den Spongien.-Zoologischer Anzeiger 13:255­ Goreau, T. E, & W. D. Hartman. 1966. Sponge: effect 167. on the form of reef corals.-Science 151:343­ Hori, M., S. Saito, Y. Z. Shin, H. Ozaki, N. Fusetani, 344. & H. Karaki. 1993. Mycalolide-B, a novel and Gray, J. E. 1867. Notes on the arrangement of sponges, specific inhibitor of actomyosin ATPase isolated 772 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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