PROC. BIOL. SOC. WASH. 104(2), 1991, pp. 209-228 new records of stylasteridae (hydrozoa: hydroida) from the galApagos and cocos islands Jill 1 >l Stephen D. Cairns Abstract.—Based on the collections ofthe Johnson-Sea-Link I submersible, new records of 14 stylasterid species are reported from the Galapagos Islands, includingone newspecies: Lepidopora concatenata. Five stylasteridspeciesare reported from Cocos Island, including two new species: Pliobothrusfistulosus and Stylaster cocosensis. These are the first records ofstylasterids from Cocos Island, three ofthe five species also shared with the Galapagos Islands. These specimens are also the first records ofthe genera Lepidopora and Pliobothrus intheeasternPacific, fromtheGalapagosandCocosIslands, respectively. Both theGalapaganandCocos Islandstylasteridfaunaeareconsideredtobederived from the western Pacific, having no affinity with the shelfand slope fauna of the American continents in the eastern Pacific. A rich fauna offourteen stylasterid spe- (1986b)andisnotrepeatedhere. Oneofthe cies, all endemic to the Galapagos Islands species previously reported, Stenohelia ro- and all based on only sixAlbatross stations busta Boschma, 1964a, is herein synony- made at the turn of the century, was re- mized with Stenohelia concinna Boschma, viewedbyCairns(1986b). Therefore, itwas 1964a, and one new species is described, anticipated that a more intensive collecting which maintains the number of species effort (27 stations in the Galapagos and 8 known from the Galapagos at 14. The ad- offCocos Island) using more efficient and ditional specimens collected by the John- selectivecollectingmethods(amannedsub- son-Sea-Link I, however, did: allow an ex- mersible) would certainly yield many more tension ofthe geographic, bathymetric, and species. Although stylasterids were sighted temperature ranges of all species; provide on every submersible dive and collected at information on the range of variation of 21 ofthe27 sitesintheGalapagos, inwhich various morphological characters; and es- all 14 previouslycollectedspecieswererep- tablish maximum colony sizes for half of resented,onlyonespecimenofonenewspe- the species. cies was collected. This surprising result StylasteridswerealsoabundantoffCocos might be explained by assuming that all of Island,beingcollectedfrom fouroftheeight the Galapagan stylasterid species are uni- stations. Since no stylasterids had previ- formlyandabundantlydistributedthrough- ously been reported from Cocos Island, the outthearchipelago, suchthatevenacursory five species reported herein are all new rec- sampling would have a high probability of ords for this island, including two new spe- collecting most ofthe resident species. This cies. hypothesis would explain both the paucity Materialandmethods.—Thenewrecords of new species and the re-collection of all onwhich this report isbased originated pri- previously known species. marily from collections ofthe R/VSeward The history ofour knowledge ofthe Ga- Johnson andJohnson-Sea-Link Isubmers- lapagosstylasteridfaunawasgivenbyCairns ible expedition to the Galapagos and Cocos 210 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON Table 1.—List oiJohnson-Sea-LinkI(JSL) stationsatwhichstylasteridswerecollected(Fig. 32). Bottomtemperature Station Latitude longitude(°W) Depth(m) Date (°C) 1911 0°32.7'S 90°07.0' 426^62 12 Nov 1986 8.3-10.3 1912 0°21.9'S 90°15.7' 713-806 14 Nov 1986 5.9-6.7 1913 1°32.7'S 90°25.8' 84-227 14 Nov 1986 13.5-14.5 1914 1°17.3'S 90°17.4' 166-172 15 Nov 1986 14.3-14.4 1915 1°17.2'S 89°48.7' 650-652 15 Nov 1986 7.4-10.8 1916 1°18.7'S 89°48.8' 545-562 16 Nov 1986 7.3-10.8 1920 1°46.6'S 89°30.8' 64-104 17 Nov 1986 1921 0°17.0'N 89°59.8' 680-720 18 Nov 1986 6.0-6.7 1922 0°23.7'N 90°26.3' 475-578 19 Nov 1986 7.3-8.9 1923 0°23.7'N 90°32.3' 348-384 19 Nov 1986 9.9-10.0 1924 0°03.9'N 90°19.2' 373^30 20 Nov 1986 1925 0°14.9'S 90°32.5' 415-447 20 Nov 1986 8.9-9.3 1927 0°10.5'S 90°53.3' 708-784 21 Nov 1986 5.6-6.5 1928 0°15.2'S 91°06.9' 720-813 22 Nov 1986 5.0-7.5 1929 0°14.7'N 91°36.5' 806 23 Nov 1986 5.7-9.3 1931 0°10.3'S 91°24.7' 441-525 24 Nov 1986 7.9-8.0 1932 0°15.1'S 9r27.7' 313-315 24 Nov 1986 10.4-12.2 1933 0°17.1'S 91°40.2' 663-788 25 Nov 1986 6.0-7.0 1935 0°15.2'S 91°27.9' 252-308 25 Nov 1986 7.8-9.0 1936 0°40.7'S 9r24.0' 182-224 26 Nov 1986 12.5-14.4 1937 0°59.8'S 91°27.1' 315-316 27 Nov 1986 11.2-12.6 1940 5°34.6'N 87°04.3' 546-631 1 Dec 1986 9.0 1942 5°34.6'N 87°04.3' 606-628 2 Dec 1986 6.7-7.1 1943 5°26.rN 87°08.0' 303-333 2 Dec 1986 11.9 1944 5°28.1'N 87°08.0' 293-576 3 Dec 1986 7.7-8.3 IslandsinNovemberandDecemberof1986. The following abbreviations are used in Stylasterids were collected at 25 of the 35 the text: submersible sites (Table 1). The use of a CDRS Charles Darwin Research Station, manned research submersible allowed se- Santa Cruz, Galapagos lective collection of deep-water inverte- H:W Height-to-width ratio ofa gastro- brates and in situ observation and photog- style rraepchoymmeofndelidvifnogr tchoeralc,ollaecntdionisofhidgeehpl-y JSL Johnson-Sea-Link I, a 4-man re- search submersible owned by the water benthic invertebrates in areas ofrug- Harbor Branch Oceanographic In- ged topography. stitution, Inc., Fort Pierce, Rorida Definitions ofthe terms used in the de- USNM Collections of the United States scriptions are reviewed and illustrated by National Museum (now housed at Cairns (1983b, 1986a). Synonymies are not the National Museum of Natural always complete; where they are not, they History), Smithsonian Institution, includeareferencetoacompletesynonymy. Washington, D.C. All type and most nontype specimens are deposited at the USNM; a synoptic collec- Zoogeography.—Cairns(1986b)reported tionisalso depositedatthe Charles Darwin that all 14 stylasterid species known from Research Station. The scanning electron the Galapagoswere endemic. Furthermore, photomicrographs were done by the author he stated that, at the generic level, the af- on a Cambridge Stereo Scan 100. finities of the Galapagos stylasterids were VOLUME 104, NUMBER 2 211 withthewestern Pacificandless sowiththe pagos Islands), 680-720 m, 1 colony and western Atlantic, not at all with the eastern SEM stub 698, USNM 84697. Pacific. The deep-water Equatorial Under- Description.—Colony uniplanar and rel- mm currentwassuggestedasapossibledispersal ativelysmall, theholotype20.4 talland mm mechanism for larvae. The collections of 30.2 broad, with a basal branch di- the Johnson-Sea-Link I expedition rein- ameter of 3.2 mm. Branching sparse, di- force this interpretation. At the species lev- chotomous, and equal, resulting in a fairly el, three Galapagan species are now known symmetrical branching pattern. Branches from Cocos Island and one, Crypthelia cy- circularin cross section, terminating in tips mm mas, isknownfromoflfNewZealand(Cairns about 1.8 in diameter; nobranchanas- 1991c). This observation, along with the tomosis. Coenosteum white and reticulate- discovery ofLepidopora in the Galapagos imbricateintexture. Coenostealstripsbroad (a widespread genus but not known else- but variable in width, ranging from 0.11 to mm where in the eastern Pacific) reinforces the 0.20 across and separated by a series hypothesis ofa western Pacific origin ofthe ofquite broad (about 66 ft wide), elliptical Galapagan stylasterid fauna. to elongate coenosteal pores and shallow Five stylasterid species are known from grooves. Platelet structure poorly defined offCocos Island, belonging to three genera: (Fig. 6), the platelets being only 18-35 n Pliobothrusfistulosus, Errina macrogastra, wide. Stylastermarenzelleri, S. galapagensis, and Gastropores uniformly distributed on 5". cocosensis. At the species level, two of branch surfaces, circularin shape(0.31-0.33 mm the five are known only from Cocos Island, in diameter), and flush with coenosteal the remaining three also occurring in the surface (no gastropore lips). Gastropore Galapagos Islands. All three genera have tubes cylindrical and deep, each bearing a widespread distributions but do not occur diffuse ringpalisade composed ofblunt, cy- in the eastern Pacific. Thus, although based lindrical elements up to 45 ju tall and 17 ix onrelativelyfewspecimens,itwouldappear indiameter. Becauseonlyonespecimenwas thattheCocosIslandfaunaisalsoprimarily available for study, a fragment was not sac- influenced from the western, not eastern, rificed to examine and figure a gastrostyle; Pacific. This is contraryto the indication of however, from gross external examination, Briggs (1974), based on shallow-water in- gastrostyles appear to be elongate and slen- vertebrates and shore fish, and Cairns der, probably with a relatively high H:W (199la),basedonahermatypicScleractinia, ratio. Dactyloporesarrangedalong sinuous, who both considered Cocos Island as an longitudinal or oblique ridges that occa- extension ofthe Panamanian Province. But sionallyanastomoseintoareticulatepattern it must be remembered that the Cocos sty- (Figs. 2, 5). Dactylopores circular to ellip- lasteridfaunaisapredominantlydeepwater tical in shape: circular dactylopores about one (e.g., 300-800 m) and therefore prob- 85 /u in diameter, elliptical ones up to 118 ably not influenced by the shallow water H in greaterdiameter. Ridges on which dac- currents from the Panamanian mainland. tylopores occur quite tall (up to 0.5 mm) andthin(25-30 wide),dactyloporecenters ij, mm New Records spaced every 0.5-0.8 along ridge. Dac- tylopores are the most prominent sections Family Stylasteridae Gray, 1847 of a ridge, the ridge connecting adjacent Lepidopora concatenata, new species pores sometimes slightly lower in height. Figs. 1-7 Colony sexually immature and therefore Records.-nololypQ-. JSL-1921, 0°17'N, sex and characteristics of ampullae un- 89°59.8'W (southwest of Genovesa, Gala- known. 212 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON Figs. 1-4. Lepidopora concatenata, holotype, USNM 84697: 1, Holotypecolony, x2.3; 2, Branchtipillus- trating tall dactylopore ridges and one gastropore, x25.5, stereo pair; 3, 4, Elongate ridges concatenating (linking)dactylopores, x56, x62, respectively. VOLUME 104, NUMBER 2 213 Figs. 5-7. Lepidoporaconcatenata, holotype, USNM 84697: 5, Branch segmentillustratingtwogastropores andridgeslinkingdactylopores, x21,stereopair;6,Weaklyimbricatecoenostealtexture, x 180; 7,Branchcross section revealing porous central core, two gastropores in cross section, and tall coenosteal ridges peripherally, x38. 214 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON Discussion.—Amongthe 25 valid species (Cairns 1991b); thus, this is the first record of Lepidopora (see Cairns 1991c), L. con- ofthe genus for the eastern Pacific. catenata is most similar to L. acrolophos Cairns, 1983a, known only from the Scotia Pliobothrusfistulosus, new species Ridge west ofSouth Georgia (659-686 m). Figs. 8-14 Both species have: similarly sized and shapedcorallaandbranchingpattern; linked Records.-UoXoXype:JSL-1940, 5°34.6'N, dactylopores; same sized gastro- and dac- 87°04.2'W (north coast of Cocos Island), tylopores; and the same bathymetric range. 546-631 m, 9°C, 1 colony (ex SEM stub USNM Lepidopora acrolophos, however, differs in 699), 84696. having a very different coenosteal texture Description.—Colony probably unipla- (reticulate-papillose), small abcauline gas- nar, the only known specimen (the holo- mm tropore lips, dactylopores that are some- type) a colony fragment 11.0 tall and mm times not linked by ridges, and in lacking 10.5 broad, consisting of 3 terminal ring palisades. branches. Branching dichotomous and ap- Although less similar to L. concatenata, parently equal; branches circularto slightly L.polystichoporaCairns, 1985(northofNew elliptical in cross section, having blunt to mm Zealand, 197-710 m) also sometimes has slightly clavate tips 1.8-2.2 in diame- linked dactylopores as well as having equal ter. Coenosteumwhiteandlinear-imbricate branching and primarily imbricate coenos- in texture. Coenosteal strips quite thin (21- teal texture. ButL. polystichopora differs in 45 Ai wide), bordered by deep, elongate slits having its dactylopores arranged in four to that are often equal to or wider than strip sevendiscrete, longitudinal rows, the ridges width (e.g., up to 55 M wide). Slits traversed uniting the dactylopores being quite low bynarrow(20-23mwide)coenostealbridges (about 0.12 mm); having small, abcauline (Fig. 10). Platelets very poorly defined but gastropore lips; and in lacking ring pali- present in a unilinear series on each strip. sades. Gastropores uniformly distributed on all In addition to L. concatenata, only one branch surfaces, circularin shape(0.37-0.39 mm other species of Lepidopora has ring pali- in diameter), and flush withcoenosteal sades: L. glabra (Pourtales, 1867), known surface. Gastroporetubescylindricalwitha onlyfromtheStraitsofFloridaat267-1 170 slightly enlarged lower section, and bent m (see Cairns 1986a). L. glabra, however, about90°,thelowersectionrunningparallel shares few other characters, differing in: to branch axis. Total length of gastropore coenosteal texture; arrangement of gastro- tube 0.9-1.0 mm. No ring palisades or ta- and dactylopores; havingunlinked dactylo- bulae present. Dactylopores also uniformly pores; and in having quite large abcauline distributed on branch surfaces and circular gastroporelips.Thecharacteroftall,slender inshape: 86-93nindiameter. Dactylopores ridgesthatconsistentlylinkitsdactylopores elevated on cylindrical mounds as high as is unique to L. concatenata. 0.2 mm, the thin walls ofthe mound 15- Etymology.—The species name concate- 20)uthick,resultinginadactyloporemound nata (Latin for "linked together") refers to diameter of0.12-0.13 mm. the dactylopores ofthis species, which are Sex of holotype probably female. Am- mm linked together by tall, thin ridges. pullaeinternal, 0.3-0.4 indiameter,and Distribution.—Species known only from communicatetocoenosteal surfacethrough type locality. Lepidopora is a wide ranging aninconspicuous,spongyefferentporeabout mm genus known from throughouttheAtlantic, 0.10 in diameter. Subantarctic, and South Pacific (New Zea- Discussion. —Four Recent species of m land region) from depths of 60-1874 Pliobothrus are known: P. tubulatus (Pour- VOLUME NUMBER 104, 2 215 Figs. 8-12. Pliobothrusfistulosus, holotype, USNM 84696: 8, Holotype colony, x7.0; 9, Branch segment illustratingone gastropore and two smallerdactylopores, x44; 10, Porous coenosteal texture with irregularly- shaped efferent pore in center, x92, stereo pair; 11, Porous coenosteal texture (dactylopore in upper right), X110; 12, Enlargementofcoenosteal texture, x 170. 216 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON USNM Figs. 13, 14. Pliobothrusfistulosus, holotype, 84696: 13, Branch tip bearing a concave, ruptured ampulla (left center), x20.5, stereo pair; 14, Longitudinal section ofa gastropore tube (above) and internal ampulla(below), x46, stereo pair. VOLUME 104, NUMBER2 217 tales, 1867) (Antilles, 419-718 m); P. sym- reportedfrom sixmorelocalities, extending metricus Pourtales, 1868 (North Atlantic, its range into the western islands ofthe Ga- 80-1600 m); P. spinosus (Hickson & En- lapagos. Considerablylargerspecimenswere gland, 1905) (Indonesia, 1089 m); and P. collected during the JSL expedition, the echinatus Cairns, 1986a (Lesser Antilles, largest {JSL-\9\A) a uniplanar colony 7.7 164-708 m). Pliobothrus fistulosus differs cmtalland9.1 cmbroad,withabasalbranch mm fromallofthesespeciesinhavingextremely diameter of7.5 (Fig. 15). The tissue is narrow coenosteal strips and much smaller a characteristic lightbrown; the corallum is ampullae.Itsdactyloporemoundsareabout white. Branches are elliptical to flattened in the same height as those ofP. symmetricus cross section; the distal, small-diameter (0.2 mm), but considerably shorter than branches being flattened in the plane ofthe those ofP. spinosus (1.2 mm), P. tubulatus colony. Although abcauline dactylopore (0.8 mm), and P. echinatus (0.4 mm). It is spinesareclearlypresentonmostbranches, further differentiated from P. symmetricus dactyloporespinesareoftenquitesmalland by having less robust branches. low, and have very short, obscure dactylo- Etymology.—The species namefistulosus tomes, thus sometimes superficially resem- (Latin for "forming a pipe," "hollow," or bling species ofLepidopora. "porous") could be interpreted in any of Distribution. —Widespread throughout three ways: forming a pipe, alluding to its Galapagos, including off: Femandina, Isa- long axial gastro- and dactylopores, which bela, south ofMarchena, northeastofSanta givethebranch averyporous cross section; Cruz, Espaiiola, and Floreana; 166-806 m; hollow, referringtothenumerousaxialgas- 5.7°_14.4°C. tro- and dactylopores and the numerous rather large coenosteal pores that penetrate Distichopora laevigranulosa Cairns, 1986 the coenosteum; and "porous," as ageneral Fig. 16 designationforthehighlypermeable nature ofthe coenosteum. Distichopora laevigranulosa Cairns, 1986b: Distribution.—Knownonlyfromthetype 6-8, figs. 3A-I. locality. Species of Pliobothrus are known fromtheNorthAtlanticand Indonesian re- New records.—JSL Stations 1911, 1 fe- m USNM gion from depths of 80-1600 (Cairns malecolony, 84704; 1914, 2 female USNM 1991b); therefore, this is the first record of colonies, 84705; 1916, 1 femalecol- USNM the genus from the eastern Pacific. ony, 84706; 1929, 2 female colo- USNM nies, 84707. Lepidotheca macropora Cairns, 1986 Discussion.—Distichoporalaevigranulosa Fig. 15 waspreviouslyknown from onlyonebranch fragment(theholotype); sixadditionalspec- Lepidotheca macropora Cairns, 1986b:4-6, imens from four more localities are record- figs. lA-G, 2A-C. edherein. Thelargest, andcomplete,colony Newrecords.—JSLStations 1914, 16 col- is 31.0 mm tall and 3.95 mm broad, with USNM mm onies, 84698; 1924, 3 colonies, a basal branch diameter of 5.2 x 6.2 USNM 84699; 1929, 5 colonies and (/5'L-1929). It has equal, sparse branching, USNM branches, 84700; 1932, 3 colonies resulting in only 10 branch tips (Fig. 16). USNM and branches, 84701; 1933, 1 Tissue light brown; corallum white. The USNM branch, 84702; 1937, 1 colony, holotype has several gastropores on its an- USNM 84703. terior face; however, the colony from JSL- — Discussion. Previouslyknownfrom only 1916 has numerous gastropores on its an- two localities in the Galapagos, 29 addi- terior face in addition to those constituting tional colonies ofL. macropora are herein the lateral edge pore row. Another point of 218 PROCEEDINGSOFTHE BIOLOGICALSOCIETYOFWASHINGTON Figs. 15-18. 15,Lepidothecamacropora,75'L-1914,USNM84698, x0.81; 16,Distichoporalaevigranulosa, JSL-1929, USNM 84707, female colony, xl.2; 17, Stylaster marenzelleri, JSL-1923, USNM 84730, male colony, xO.8; 18, Stylastercocosensis, holotype, USNM 84734, x1.3. variationisthatoneofthedactyloporerows Distribution.—OSKoca. Redonda, Santa ofthe specimen fromJSL-1911 has almost Cruz, Espanola, and Floreana; 166-806 m; completely degenerated, the opposite row 5.7°-14.4°C. Species of Distichopora are appearingtobethe onlyfunctionaldactylo- common in the Indo-West Pacific (Bosch- pores. The holotype, as well as all six spec- ma 1959) and western Atlantic (Cairns imensreportedhere, are maturefemalecol- 1983a), but are conspicuously absent from onies, thus male ampullae remain un- the Antarctic-Subantarctic region and east- known. em Atlantic. Distichopora laevigranulosa is