SCI. MAR., 57(4): 381-393 SCIENTIA MARINA 1993 RECENT ADVANCES IN ECOLOGY AND SYSTEMATICS OF SPONGES. M.J. UR1ZURIZ and K. RUTZLER (eds.).(eds.). The genus Terpios (Suberitidae)(Suheritidae) and new species in the «Lobiceps» complexcomplex** KLAUS RUTZLER and KATHLEEN P. SMITH Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution,Institution, Washington,Washington, D.C.DC. 20560, U.S.A. SUMMARY: The genus Terpios, first described from thethe Atlantic Ocean (West(West Indies)Indies) almostalmost 130 yearsyears ago,ago, isis redefined on the basis of structural studies of tylostyles and data on procaryotic symbionts.symbionts. ItsIts relationrelation toto similarsimilar generagenera in the family Suberitidae is reviewed by comparing new findings with traditionallytraditionally used characters, suchsuch asas bodybody shape,shape, skeleton arrangement, and spicule size. Two new species, T. manglaris and 7".T. belindae, areare described fromfrom shallow-shallow­ water habitats in the Caribbean Sea. They resemble Suberites lobicepslobiceps Schmidt, a poorly knownknown spongesponge fromfrom FloridaFlorida that has not been found since its first description inin 1870. Key words: Porifera, bacteria, symbiosis, Terpios, new species, Caribbean. INTRODUCTION Because of its shape and striking color, Terpios fugax is very distinctive, and the genus Terpios has In their pioneering study of Caribbean sponges, remained valid for almost 130 years, although it was DUCHASSAING and MICHELOTTI (1864:97) introduced not universally adopted. VON LENDENFELD (1897: the genus Terpios for thinly encrusting (membrani-(membrani­ 132), for example, who found the type species in the form) sponges in which the spicules are arranged hap-hap­ Adriatic, redescribedredescribedit it under the name Suberites fu-fu­ hazardly except for some organization into fan-fan­ gax.gax, However, he failed to recognize the taxonomic shaped bundles. They described nine species in four value of spicule-size classes, as they occur in Suberites live-color groups. Their diagnosis may have led Vos-Vos­ proper, and of the peculiar shape of Terpios tylos-tylos­ MAER (1887:359) to place T. fugaxfugax in this genus and tyles. The same was true of his colleague TOPSENT DE LAUBENFELS (1936a:(1936a:152) 152) to select it as the geno-geno­ (1900:(1900:192f),192f), who argued that Terpios is distinguished type,type. T. fugax not only conforms to the original defi-defi­ from Suberites by encrusting (rather than massive) nition but is the only encrusting species of the re-re­ habit, gelatin-soft consistency, a smooth surface with-with­ maining three of the original nine that are still out spicule reinforcement of the ectosome, and regu-regu­ recognizable (VAN SOEST, etai,et at" 1983:204). The other lar and loose skeleton structure. TOPSENT (1900:194)(1900: 194) two are T. aurantiaca, definitely a massive species, pointed out that the tylostyles of T. fugaxfugax are non-non­ and T. janiae,janiae, which is a Dysidea. fusiform, as the shaft progressively thins to a sharp point, and their heads are quite variable, from glob-glob­ * Received February 10, 1993. Accepted June 2, 1993. ular with tapered top to depressed, some being tri- NEWS SPECIES OF TERPIOS 381 lobed and some having annular swellingsswellings inin thethe neckneck alive and with thethe help ofof phase-contrastphase-contrast microsco­microsco- region. These observations werewere expandedexpanded uponupon byby py. Museum material was fixedfixed inin 1010 %% formalin­formalin- DE LAUBENFELS (1936a:(1936a: 152),152), whowho claimedclaimed thatthat seawater and preserved inin 8080 %% ethanolethanol afterafter 2424 h.h. Terpios is "characterized by thethe quadrilobatequadrilobate formform It was deposited inin thethe collectioncollection ofof thethe NationalNational of the heads of spicules thatthat otherwiseotherwise wouldwould bebe re­re- Museum of Natural History, SmithsonianSmithsonian InstitutionInstitution garded as tylostyles." Subsequently,Subsequently, DEDE LAUBENFELSLAUBENFELS (USNM). (1950:(1950:103)103) was struck by thethe uniqueunique morphologymorphology ofof The skeletal structure was examinedexamined inin sectionssections Terpios tylostyles. InIn particular, hehe consideredconsidered thethe 100 ~m\im thick that were prepared byby grindinggrinding andand pol­pol- relative size of thethe head distinctdistinct amongamong thethe suberitids,suberitids, ishing sponge tissue samplessamples embeddedembedded withwith epoxyepoxy for it was "nearly double oror quitequite doubledouble thethe diame­diame- resin (as described by ROTZLER,RUTZLER, 1978).1978). SpiculesSpicules werewere ter of the spicule shaft."shaft." HeHe alsoalso observedobserved thatthat "in"in isolated and cleaned by boilingboiling inin concentratedconcentrated young sponges" thethe head isis distinctlydistinctly lobate,lobate, "with"with nitricacid and washing inin demineralizeddemineralized waterwater andand indications that thethe laterallateral growthgrowth ofof thisthis headhead hashas absolute alcohol. Measurements areare basedbased onon 2525 ty­ty- (primitively) arisen by polyactinal branching.branching. ItIt maymay lostyles selected at random.random. ToTo revealreveal internalinternal struc­struc- represent a pentactinal spiculespicule withwith fourfour cladsdads inin oneone ture~ure and enhance thethe axialaxial canal,canal, somesome spiculespicule sam­sam- plane, and a very longlong rhabd."rhabd." HisHis interpretationinterpretation ofof ples were concentrated by centrifugationcentrifugation inin epoxyepoxy Terpios appears toto have been biasedbiased byby hishis earlierearlier resin, ground and polished (Carborundum(Carborundum paperpaper andand examinations of encrusting specimensspecimens withwith distinctlydistinctly aluminum oxide, toto 0.3 ~ri m), andand etchedetched byby exposureexposure lobed heads because thethe ensuingensuing descriptiondescription andand dis­dis- to dilute hydrofluoric acidacid (6.5(6.5 %% inin distilleddistilled water)water) cussion of "T. fugax" does notnot mentionmention thesethese charac­charac- for 15 sec. Scanning electronelectron micrographsmicrographs (SEM)(SEM) teristics at all. De Laubenfels was,was, inin fact,fact, describingdescribing were made of spicules andand etchedetched sectionssections usingusing aa T. aurantiaca DUCHASSAING andand Michelotti,Michelotti, notnot thethe Cambridge Stereoscan 100100 microscopemicroscope atat 1,000­1,000- massive growth form or stagestage ofof T.T. fugax,fugax, asas hehe pre­pre- 3,000 xxmagnification.magnification. sumed. For transmission electron microscopymicroscopy (TEM),(TEM), It is not uncommon forfor thethe tylostylestylostyles ofof crustosecrustose material was fixed inin 3.5% glutaraldehydeglutaraldehyde inin 0.10.1 MM sponges to have conspicuously lobedlobed heads.heads. ThisThis fea­fea- phosphate buffer with thethe additionaddition 0.450.45 MM sucrosesucrose ture is seldom mentioned inin thethe literatureliterature becausebecause itit (90 min at 29"29° C), postfixed inin 11 %% osmicosmic acidacid inin thethe is usually seen inin small,small, fragmentary,fragmentary, oror isolatedisolated sam­sam- same buffer mixture (60(60 min atat 4°4" C).C). SectionsSections werewere ples. One exception isis SCHMIDT'SSCHMIDT'S descriptiondescription andand il­il- stained in saturated (5(5 %)%) alcoholicalcoholic uranyluranyl acetateacetate lustration (1870:47; pI.pi. 5,5, fig.fig. 5)5) ofof anan unusualunusual spongesponge with 0.25 % leadlead citrate andand viewedviewed andand photo­photo- crust from Florida, which hehe namednamed SuberitesSuberites lobi­lobi- graphed through a leolJeol 12001200 EXEX electronelectron microscopemicroscope ceps. A specimen with similarsimilar spiculesspicules fromfrom thethe GulfGulf (2,000-12,000 x primary magnification). LightLight mi­mi- of Mexico (TOPSENT, 1920:30)1920:30) waswas identifiedidentified (but(but notnot croscope observations ofof histologyhistology andand skeletonskeleton described) by SCHMIDT 1880:77,1880:77, butbut thisthis speciesspecies hashas structure were made on thethe samesame materialmaterial sectionedsectioned 11 not been found or discussed sincesince thesethese earlyearly reportsreports urn~m thick, or ground and polished toto aa thicknessthickness ofof 5050 and remains obscure; DE LAUBENFELSLAUBENFELS (1950:107)(1950:107) urn,~m, both stained by methylene blue.blue. dropped it in synonymy with T.T. fugax.fugax. The question of validity ofof thethe genusgenus TerpiosTerpios waswas reopened recently inin aa discussion ofof stronglystrongly compe­compe- RESULTS titive Pacific reef speciesspecies ofof TerpiosTerpios withwith distinctlydistinctly lobed tylostyle heads (ROTZLER(RUTZLER andand MUZIK,MUZIK, 1993).1993). 1. Superspecific Characterizations The question could not be addressedaddressed withoutwithout revie­revie- wing T. fugax, thethe genotype, andand thethe statusstatus ofof Suber­Suber- The following diagnoses were compiledcompiled fromfrom vari­vari- ites lobiceps and other closelyclosely relatedrelated speciesspecies withwith un­un- ous sources, primarily TOPSENT (1900),(1900), DEDE LAUBEN­LAUBEN- usual tylostyle heads, includingincluding thethe twotwo newnew onesones FELS (1936a), and LEVI (1973),(1973), andand supplementedsupplemented byby from the Caribbean described inin thisthis report.report. our own observations. CommentsComments onon andand descrip­descrip- tions of examples forfor different generagenera areare basedbased pri­pri- marily on tropical western AtlanticAtlantic