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East Pacific Mexican Tethya (Porifera: Demospongiae) with descriptions of five new species PDF

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OFWASHINGTON 114(3):794-82L 200L East Pacific Mexican Tethya (Porifera: Demospongiae) with descriptions of five new species Michele Sara, Patricia Gomez, and Antonio Sara (MS, AS) Dipartimento per lo Studio del Territorio e delle sue Risorse, Universita di Genova, Corso Europa 26, 16132 Genova, Italy (PG) Instituto de Ciencias del Mar y Limnologia, U. N. A. M. Circuito Exterior S/N, Ciudad Universitaria, C.P. 04510, Mexico, D.E — Abstract. Seven species of Tethyidae (Porifera: Demospongiae: Hadrom- erida) inhabiting the Pacific waters ofMexico, including the GulfofCaHfornia, were studied. Five of them, Tethya ensis, T. mexicana, T. ovum, T. paroxeata, and T. socius are new species. The remaining two, T. taboga De Laubenfels and T. californiana De Laubenfels, are new records for the area. This is the first published report on spe- sis group or if it should be considered as cies of the genus Tethya (Porifera: Demos- representative of a new genus ofTethyidae. pongiae) coming from the Pacific coast of Mexico. A total of 242 collected Tethya Materials and Methods specimens yielded five new species andtwo other species recorded for the first time. Samples of Tethya (242 individuals) rep- Tethya taboga (De Laubenfels) was known resent part of several collections obtained only for the Pacific coast of Panama, and along the Pacific coast ofMexico, including T. californiana De Laubenfels, 1932, rede- the Gulf of California at different depths scribed by Sara & Corriero (1993), is re- (Table 1, Fig. 1). Sponges were collected with SCUBA diving and with a trawl corded here for a more southern site in aboard the R/V El Puma. Thirty-eight Mexico. Only four species of Tethya were previously known for the whole Pacific trawls, covering almost all the Gulf of Cal- ifornia from 31°15'9"N, 114°21'7"W to coast of America: T. californiana De Lau- m 20°49'N, 105°42'W, from 22 to 120 benfels from California, T. taboga (De Lau- depth, were performed in March, July, Au- benfels) from Panama, T. papillosa Thiele gust, and October of 1985. Several dives off from Calbuco (Chile) and T. sarai Des- the coast of Mazatlan took place from queyroux-Faundez & van Soest, 1997 from 23°15'N, 106°29'W to 23°11'30"N, 106° the Galapagos Islands. This small number 25'W in May 1981 and June 1987 at a of American Pacific species, now amount- depth of to 15 m. Trawls and dives from ing to nine, is probably due both to a real Guerrero were performed in February scarcity of species and to the poverty ofre- (Winter) and April (Spring) of 1982, in the cords from this area. Donatia multifida continental shelf, from the coastline to the Carter, 1882 from Acapulco (Mexico), ex- beginning of the continental slope, at 200 amined by Sara (1994) on spicular slides of m depth (102°15'N, 98°W), including the the British Museum of Natural History Balsas river delta. Sponges, once aboard, (BMNH) is currently regarded as incertae were preserved first in 95% alcohol and sedis, given the scarcity of the material. then in 70%. Techniques to analyze internal Moreover, it is impossible to know if it be- structures follow Sara (1992), and scanning longs to a species ofthe Tethya seychellen- electron microscopy (SEM) follow Gomez VOLUME 114, NUMBER 3 795 (1998). Spicular data reported are based on 50 measurements from each spicular trait. Main and auxiliary megascleres, generally, ^ strongyloxeas, were distinguished by a con- ventional 1000 |xm length figure. For two P3 -O ^ Z 13 species, a third category of "sword" stron- iri ^ Z ^Z^ -^ cglyaltouxreeasiswatrsadciotnisoindaellryed.baMsiecdrasotneronbsoemrevna-- CL, (^<iUi (-^NH ('SN^; m^^ (/Cc:O) tions taken at the light microscopic level. Yet, the difference between oxyasters, strongylasters and tylasters (Sara 1994) is only partially due to the ray shape. It de- pends largely, as shown by SEM micro- graphs, on the distribution and strength of the spines. The holotypes are deposited in u 00 ID the Natural History Museum of Genoa (It- rmo^ ^ aly) (MSNG), some paratypes are deposited oa >-< in m•^ r^o 055 in Laboratorio de Ecologia de Bentos ICML collection of Mazatlan (Mexico). — Study area. The Gulf of California be- longs to the subtropical regime with marked fluctuations in climatic conditions all year z = long. It is positioned between two dry con- ;O.n. QO 0"0^ i3s tinental environments that cause wide rang- )Ph ^ '^ 00 '^ ^ (N-H(NH^(N^mM es in temperature, low humidity, and high evaporation rate. The yearly mean temper- ature value at the surrounding coast is 24°- 26°C. Rainfall is abundant at the east coast of the Gulf as well as in the south, whereas at the west coast rainfall is lower. Maxi- z C mum rainfall is from June to October, be- coming dry at winter and at the beginning u m 0< of spring. This brings about coastal up- welling along the western side in summer and at the eastern side in winter (INEGI 1984, Molina-Cruz 1986). Mazatlan is lo- cated at the mouth ofthe GulfofCalifornia, in a tropical and subtropical semi-humid rt 00 climate, bathed by the Gulf of California 73. <^ current and causing different water changes ^ M O ^ vo ro c/:i according to the nearby inlets, islands, and breakwaters. Yearly mean temperature is from 25°C to 28°C, yearly mean rainfall mm U reaches 850 (Alvarez-Leon 1980). Guerrero has a tropical semi-humidclimate, 0«) ? 7033 3^ 5 S'^ yearly mean temperature of 27.5°C, yearly c 03 G mean rainfall 1117 mm, and a permanent o ^&^ 6 >. influx of freshwater in Petacalco Bay that Uo Qu Uo H(U OX m03 796 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ 115° 110° 105° 100° 95° 90° >^j S;\ .:^S \' P. Libettad . T.sodiis ^^^fi-'^^-^^M C.SanMTpu^^QN T.cahformam T^<^ - . ^ ^^boleda paroxeata mexicana \ \ .J Mexico ( Gulfof IVIexico^ I) ^.\Mazatlan "I \y T.taboga 1 ^ \ East Pacific "'^v R. Balsas -- I B. Petacaico '<^^^ T.mexicana ''^--^ T. ovum P. MaldonadoS^-.<._^^ En T.mexicana ^ 115° 110° ^1^0^5^° T.ensis ^10^0„° 95° \^ 90° Fig. 1. Map ofcollecting areas and distribution of Tethya species: Puerto Libertad, Cabo San Miguel and Punta Arboleda inside the Gulf of California, Mazatlan at the mouth, and Petacaico Bay as well as Punta Maldonado at Guerrero, southern Mexico, tropical East Pacific. comes from the Balsas river (Amezcua-Lin- oped cortex distinct from the choanosome, ares 1996). megascleres bundles radiate from the center The study area, except the Gulf of Cali- of the sponge ending in tubercles on the fornia, represents the transition zone be- surface. Main and auxiliary megascleresare tween the mixture from the North Califor- usually strongyloxeas, auxiliary may be nian current and the South Peru current also styles, megasters are spherasters or ox- with the North equatorial counter-current, it yspherasters, micrasters are tylasters, stron- marks the limits between the Californian gylasters (chiasters) or oxyasters; these two Province with the Panamic one, now sub- are distributed in the cortex as well as in divided in Cortez Province, Mexican Prov- the choanosome (Sara 1994). ince, and Panamic Province (Hendrickx 1995). Tethya ensis, new species Figs. 2A, B, 3, 4, 15A, Table 2 Systematics — MSNG Material examined. Holotype: Order Hadromerida Topsent, 1900 50191; paratype: LEB-ICML 255; 46 spec- Family Tethyidae Gray, 1867 imens Punta Maldonado (Guerrero), Feb m Genus Te—thya Lamarck, 1815 1982, 45 dep—th. Type species. Tethya aurantiwn Pallas, Description. The type has an elongated 1766. hemispherical shape, 1.5 by 2.5 cm, with — Diagnosis. Spherical or subspherical the irregular flattened basis covered by ro- body not supported by a stalk, well devel- bust filamentous stolons. On the whole, the VOLUME 114, NUMBER 3 797 shape is elongated hemispherical (Fig. 2A, rays. Similar in the cortex and in the choan- B), 0.8-2.6 by 0.7-3 cm. Two specimens osome. — were attached to the next Tethya species de- Etymology. From Latin ensis = sword, scribed herein. Color: not recorded in vivo, in reference to the sword-like appearance of creamy white in ethanol. Consistency the short fusiform strongyloxeas; here used slightly compressible. Surface sparsely and as a noun in—apposition. irregularly tuberculate, partially smooth. Remarks. Tethya ensis is characterized The best distinguished tubercles are finely by a peculiar shape and the occurrence in mm mm hispid, flattened, 1—2 in diameter, 1 the cortex of the sword shortened fusiform in height (Fig. 2A), sharp and stout fila- strongyloxeas. The species is akin to a mentous stolons on the basis and the edges group of Mexican species subsequently de- of the sponges are 2 cm long. Some tuber- scribed herein and to T. californiana De & cles on the edge of the sponges produce Laubenfels, 1932 (Sara Corriero 1993). mm also 3 elongated buds. Cortex thick- Tethya ensis differs from T. californiana in ness, without tubercles, 0.5-1 mm. body shape and cortical structure and in — Skeleton. Megasclere bundles radiate some significant spicular traits: the mega- from center to cortex sometimes in a coiled scleres are strongyloxeas instead of aniso- way as in the holotype. Bundles, 245-325 strongyles (but a Mexican population of T. |jLm in diameter, ending in compact cortical californiana subsequently described herein fans without subdivisions (Fig. 3A). Me- also has strongyloxeas); the occurrence of gasters regularly distributed in the middle peculiar sword shortened strongyloxeas and, more densely, in the lower cortex, which are absent in T. californiana; and the forming a belt around the choanosome. larger size and the lower R/C of its megas- Some smaller megasters in the outer choan- ters that are spherasters instead of oxy- osome (Fig. 15A). spherasters. — Spicules. Table 2 summarizes measure- ments taken from 5 specimens. Megascleres Tethya mexicana, new species (Figs. 3B, D, 4A) are: main strongyloxeas Figs. 2C-G, 5-7, 15C, Table 3 (maximum size 2312 by 47 |xm), peculiar — MSNG shortened fusiform strongyloxeas ("sword" Material examined. Holotype: like) in the cortical fans and among the sub- 50192; paratype: LEB-ICML 256; 75 spec- cortical interstitial megascleres, 225-965 imens from Punta Maldonado (Guerrero), |jLm, with a slender head 4-8 fxm thick and Feb 1982 and 10 specimens from Petacalco a thickness of generally 15-25 ixm, in the Bay (Guerrero), Apr 1982, 45 m depth. Ad- central and distal parts of the spicule and hered to bryozoans and calcareous frag- auxiliary megascleres, from slender stron- ments on sand and sand-mud-clay bottom; gyloxeas to thin styles. 43 specimens from Punta Arboleda (Gulfof m Megasters (Figs. 3D, 4D): Spherasters Cahfornia), Jul 1985, 22 depth. — heterogeneous in size and shape, with some Description. The type is irregularly el- slight differences among the specimens and lipsoidal, cushion-like, with two opposite a main size range 50-90 |xm (maximum di- faces 3 by 2 cm, both coveredby irregularly mm ameter 115 |jLm). The R/C main range is rounded tubercles, 1—3 in diameter, mm 0.5-0.9, sometimes in the larger spicules it 0.5-2 in height. The thickness of the is 1-1.1. Ray number: 14-16. cushion is 1.5 cm. Long filamentous rooting Micrasters (Figs. 3C, 4B, C): Mainly processes are at one side of the faces, sug- strongylasters but variable from slightly gesting that in life the specimen was erect knobbed tylasters to slightly tylote oxyas- 2 cm high and 3 cm broad. Some elongated ters 9-13 |xm in diameter (minimum 5.9 tubercles with spear like buds are on the and maximum 16.5 |xm) with 12 thin spiny top. On the whole, body variable in shape. 798 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. A, B: Tethya ensis, new species. A, Holotype; B, Paratype. C-G: Tethya mexicana, new species. C- E, Three specimens from Gulf of California; F, Holotype from Guerrero; G, Transverse section of another specimen from Guerrero showing coiled skeleton and a central nucleus. H-J: Tethya paroxeata, new species. H, Holotype lateral view; I, Holotype, transverse section showing a coiled skeleton radiating from an asym- metrical center; J, Another specimen, lateral view. Scale bars = 1 cm. VOLUME 114, NUMBER 3 799 Fig. 3. Tethya ensis, new species. Light microscopic photographs. A, Skeletal structure of tubercle; B, Fusiform strongyloxeas and some megasters; C, Micrasters; D, Megasters and megasclere endings. Scale bars = 100 (xm. ^*1 ' ^< 1 1 800 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with irregularly hemispherical, ellipsoidal s£c ^ vId»Tdo) ^t>_r;^i ^^(N Ovr^~nN) e^r-n^ oaGr)m.eptCoeolrlyohaernddorra0al.n5gs-ep2e.wc5hicmeenmnsai,lniv0hee.,i8gd-ih4rttyc(Fmwihgi.itn2edCoi--r si5 ^O7N ("T7) 'q-' od'--' '0T0" brownish in ethanol. Consistency firm, u-i 00 ON slightly compressible. Surface tuberculate, with irregularly rounded or roughly polyg- o— 0d0 ^—^ OdN 0d0 onal tubercles 1-3 mm in diameter, 0.5-2 Us ro^ VdO dr^ ^d NdO mm in height irregularly spaced and un- 4d d4 dlA d4 d4 evenly developed in Guerrero specimens, u. si sometimes flattened, sometimes papillose, MoD >n oVO oNO covered by sand in the Gulf of California oae> ^ -r^^ 'r-^I 0"N-0Of" r^ specimens, others are scarcely visible with 'q 00 00 r^ r^ N4ip a surface covered by several stalked buds 2 mr^ r(N^ (-N^ ^NO en cm long, and several with visible oscules. Specimens from Guerrero, as in the type, m •^ (0(NN0 CCNN o(N c^<7-) C6N have on the basis and edges of the body CTvl^OOir^;- 4-7 flattened or filamentous rooting processes rn rn mm mm 10-16 long by 1-10 wide. Cortex g- (Fig. 5C, D), including tubercles, 2-4 mm s o O o (N IT) <3 Ct^^ 0O0n o(\^ OOnn 0CT0N thick. Spicular tufts hispidating the tuber- _Jcf^all .rV0rt—^0o^I Or^01m-<0H^ '6omIIsTTt))- 6<1UN(—r-NO)) mV6Om1--S)7 tcwleiextshasnaudpdpi3oa5rm0te-etd9e5rb0yof|bJ3uLm0nd0iln-e4st8h0eofcjshxtomraoinnnogstyholemoxece.oars- — Skeleton. The megasclere tracts branch vO ^ CM en at different levels of the choanosome (Fig. «) ^J= (r—N^ (rN^ (ITN) (NNO (>No 5B). The interstices among the tracts in the _"SM>o. '^ (1^N^ 4^^ 6'""' 6' ' 6^^ ubupnpdelrepsarotfofautxhielciahroyanmoesgaosmcelearreesf.illMeedgwaist-h C O tc OOn o O t^ >n ters are placed densely in the central and (N 00 r^ NO '5a s ^m CI5> mt^ e1n^ (ONN lower parts ofthe cortex that shows several Jc '-N^J- l4O aT6|\- ^uHo ^t^ lacunes (Fig. 15C), although it is less la- cfs ^m t^ ^r^ >(nN cunar than Tethya californiana. In some en <N (N CN specimens the megasclere tracts are coiled v•~*~ ^uo \mD CUIO NenO and sometimes they depart from a centmraml s^: 6m 4 ^ On r^ nucleus (Fig. 2G). The nucleus, about5 5 in (0N0 ^(N6 (0^N0 6CN in diameter, is made up of an irregular net- ^^ work of small styles and subtylostyles (Fig. 2cC3 £W) ^^(^i^H^^Nn mCT(m(-NNNH C0V0(oN0O0N OC4CoNNn O^(01—Nn\ 5mAe)nS.ptiscutlaekse.n—fTraobmleth3reseusmpmeacriimzeenssmoefasGuurere-- hsJ ^V^—O ^^'—D i•—n ^t^^ ^Qi^n rero and three specimens from the Gulf of O iori C^N Q oIT) O o m California. Megascleres (Figs. 6A, 7B) are ^^^ ' ' ^~* ' ' ^^ main strongyloxeas and auxiliary (cortical) 'aa? strongyloxeas withintermediates amongthe s0) £S" two categories. Maximal length is 2130 (xm yD, "Xo and thickness 41 [xm. The nucleus of some !Z) -H (N r^ 'vt- in specimens presents heterogeneous styles VOLUME NUMBER 114, 3 801 Fig. 4. Tethya ensis, new species. SEM micrographs. A, Megascleres andsomemegasters;B,Tylaster(more frequent); C, Chiaster (less frequent); D, Spheraster. and subtylostyles frequently somewhat Arboleda (Gulf of California) show sharp curved, 300-800 by 10-18 lam. differences. The main size range is 60—75 Megasters (Figs. 6A, 7C) are spherasters- |jLm and the main R/C range respectively, oxyspherasters: heterogeneous in size and 0.5-0.7 and 0.7-0.9. Ray number 16-18, shape, they vary also among the species. frequently bent, sometimes twisted or The main size range is 70-120 |xm (maxi- forked. mum 128 |jLm) and R/C range 0.7-1.2 Micrasters: Mainly strongylasters (chias- (maximum 1.4). Two specimens from Punta ters) but variablefrom slendertylasters with 802 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Tethya mexicana, new species. Details of structure. A, Nucleus skeleton; B, Branching of a mega- sclere bundle in the choanosome; C, Cortical structure; D, Cortical structure with megasclere fans. Scale bars = 100 M,m. ' 1 ' 1 ' VOLUME 114, NUMBER 3 803 slightly knobbed rays to slightly tylote ox- yasters. Sometimes with a more or less de- in uo IT) q q veloped center. They are in the cortex and v^ u-i u-i iri i/-i uus ^ VD 'O rn rn in the choanosome alike, generally 8-13 §Q ^^ ^H -^ ^^ -^ 0r^0 |xm in diameter (maximum 15.5 and mini- 0'0d 060 l0o0 q00 q00 l(NO mum 5.2), with 8-14, generally 12, thin rays (Figs. 6B, 7A, D). — Etymology. Named after its origin, the (N -^ ^ o ^ m Mexican wat—ers. u 0o0 6 OCT\ VOD or^ oo\ Remarks. Tethya mexicana is charac- Dd Wo-1 ol~~ ^O Oro IoT) voO terized by a variable, generally depressed, body shape, uneven tuberculate cortex, sometimes with long stolons as in T. ensis, o sometimes with buds, large and variable in 0o0 oo rO<n~i 0(0N size, spherasters-oxyspherasters with rays O On frequently bent, micrasters similar in the 00 00 o 0\ r<-) cortex and in the choanosome varying from a slightly tylote oxyaster to strongylaster shape or true slender tylasters. There is some intraspecific variability in spiculation, ^I'O|u->|(NIr^iIw-) with differences especially among the spec- ^ iri iri Tt ^ ^ imens ofGuerrero and PuntaArboleda. One of the Arboleda specimens (6) is character- ized by smaller megascleres and micrasters ^ O O 1^ ON 0>00 ON 0t^0 OONn (Table 3). The general structure and the On On On ON On On "or>msrf^\i-l|1cvot^on||IoNnTOo)|hIrirt^o^|l><olrN^illO4oooo esnpsiicsulbaurttraalistos hsaovmeesreevmeararlkaabfflienidtiifefsewrientchesT:. (^ IrT<)n 'CvOO -(*n cInT|) O-*n the body shape even ifvariable, is not elon- gated hemispherical as in T. ensis; the skel- etal structure shows a branching of megas- clere tracts and interstitial strongyloxeas CI NO On ON 00 '='1 which are absent in T. ensis; the sword-like (N (N (N (N CN cnI 4 4 00 NO On 6 strongyloxeas of T. ensis are lacking in T. mexicana; the megasters (spherasters-ox- yspherasters) are considerably larger in T. O O O IoT) Om mexicana and with a greater R/C than in T. o(N (or-N~ 0C^-N0l (N Ci-NH COCNN ensis. Tethya mexicana is also similar to T. O<6c^HnDn ((Q>o'—NNo 0CC^'—TN0) 4NQI'o—TO) 0"Qo'i—n0^ 6'-mCo—NNf tcraaliitsf:ortnhieanbaodbyutsdhiafpfeersisinnotsosmpeheriimcpaolr,thaanst (ON o CoT) o o o uneven tubercles and different cortical structure; the denser distribution of megas- ters in the lower cortex; the presence of '5" strongyloxea instead of anisostrongyle type "Ks^&(oU O^032SO)i '31Euo) "sQu3o ^tX^"T<;io3 Xr"•5<uI^oaii Ip<x"TOs>o3^ olslfoiwmgehertglarysacytlynelrouteme;beoaxrgyroaesfattietesrrmsteiygzpeae,stloeofrwsei;trstRhm/eiCcmroaarnsed- ^ (N ro -sf >n NO ters; and the lack of spherules.

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