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Contribution to the knowledge of the family Caecidae: 16. Revision of the Caecidae of Easter Island (Chile) (Caenogastropoda: Rissooidea Gray J. E.,1847) PDF

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© SociedadEspañola deMalacología Iberus, 23 (1): 49-65, 2005 Contribution to the knowledge ofthe family Caecidae: 16. Revisión of the Caecidae of Easter Island (Chile) (Caenogastropoda: Rissooidea Gray E., 1847) J. Contribución al conocimiento de la familia Caecidae: 16. Revisión de los Caecidae de la Isla de Pascua (Chile) (Caenogastropoda: Rissooidea E. Gray, 1847) J. Bret RAINES* and Mauro PIZZINI** Recibido el26-X-2004. Aceptadoel11-11-2005 ABSTRACT With the exception oftwo species, members ofthefamilyCaecidae from Easter Island have been previously neglected. Based on type and additional material, a revisión ofthe species known to date from Easter Is. is herein proposed, with the description of 5 new species: Caecum rehderi spec. nov., C. heterochromum spec. nov., C. pascuanum spec. nov., C. rapanuiense spec. nov., C. campanulatum spec. nov. RESUMEN Con la excepción de dos especies, los miembros de la familia Caecidae de la Isla de Pascua han sido previamente desatendido. Basado en el tipoyen el material adicional, se propone aquí dentro, una revisión de las especies conocidas hasta la fecha de la Isla de Pascua, y incluye la descripción de 5 nuevas especies: C. rehderi esp. n., C. heterochro- mum esp. n., C. pascuanum esp. n., C. rapanuiense esp. n., C. campanulatum esp. n. KEY WORDS: Mollusca, Caenogastropoda, Rissooidea, Caecidae, taxonomy, new species, Easter Island, WesternPacific. PALABRAS CLAVE: Mollusca, Caenogastropoda, Rissooidea, Caecidae, taxonomía, nuevas especies, Isla de Pascua, PacíficoOccidental. INTRODUCTION Easter Island (Fig. 1) is found inato- graphical province from the Polynesian tally isolated position approximately province. The submarine seascape fea- 3,500KmfromthecoastofChile. Froma tures widely scattered coráis affixed to zoogeographicalpoint ofview, Easter Is. the rugged volcanic substrate. The de- is inaverypeculiar áreawithintheeast- paupératebenthiccommunityemploysa ern Indo-Pacific región, and to thepoint varietyofadaptivestrategiesforsurvival that, in 1965, Schilder proposed the Ra- in an environment stressed by waves, panuian province as a sepárate biogeo- currents and the absence ofmineral nu- * ResearchAssociate, Natural HistoryMuseum ofLosAngelesCounty, P.O. Box612Victorville, CA92393 USA. e-mail: [email protected] **LargodellaCaffarelletta, 6, 00179 ROME (Italy). e-mail: [email protected] 49 Iberus, 23 (1), 2005 trients. Mostofthe coráis and otherbot- been completed during the last ten tominvertebratesaretypicaloftheIndo- years. With regard to Caecidae, during Pacificreefs,butreefshavenotformed. the three trips to Easter Island, the The circulation pattern, and espe- authors of the present work collected cially the marked upwelling between more than 350 specimens of this family. the South Pacific and Mentor currents, Rehder mentions only two species: contributes to the isolation of the área. Caecum cf. solitarium Oliver, 1915, and C. In terms of marine ecosystems, this in- amydroglyptum Rehder, 1980. In all other sularityhas alsoproduced ahighdegree publications, which we have reviewed, ofradiationinmany groups, and partic- Caecidae are listed simply as 'Caecum ularly in the caenogastropods. Thus the species'. No other descriptions or illus- fauna can essentially be described as trationswereprovided. typical Pacific fauna with a relatively So we present herein a revisión of highnumber ofendemic species. The is- the species known to date from Easter land's marine benthic fauna is generally Is., based on type and additional mater- characterised by a high degree of ial, with the description of five new species diversity and a low abundance, species. and this is true for both hard and soft bottoms. The meiobenthic family Caeci- Abbreviationsused: daealsofollowsthispattern. The Caecidae of the Easter Island AMS: Australian Museum Sydney, have been scarcely studied in the past, Sydney (Australia) with the exception of the work of CM: Canterbury Museum, Christchurch Rehder (1980). Past surveys have (NewZealand) reported endemicity rates within the LACM: Natural HistoryMuseum ofLos molluscan fauna ranging from 37% to AngelesCounty,LosAngeles(U.S.A.) 42% (Rehder, 1980, DiSalvo, Randall MNHN: Museum National d'Histoire and Cea, 1988, Raines, 2002). Rehder Naturelle,Paris (France) also indicates that some species appear MPR: MauroPizzinicollection. to have a dual relationship with certain NHML: Natural History Museum, species from Hawaii, as well as species London(U.K.) from the Kermadec Islands. In prepara- NMNZ: Museum of New Zealand Te tion for his 1980 publication, Rehder Papa Tongarewa, Wellington (New reviewed all previous studies and expe- Zealand) ditions. In addition to examining all the NSMT: National Science Museum, Easter Is. material in US museums, he Tokyo (Japan) also examined the full store of material USNM: National Museum of Natural housed in the Museo National de Histo- History,WashingtonD.C. (U.S.A.) ria Natural in Santiago, Chile. In all, WAM: Western Australian Museum, Rehder examined over 7,000 specimens, Perth(Australia) ofwhich 3480 were collected during his triptotheislandin 1974. Terminology: In the mid 1980's, DiSalvo and his teamconductedacomprehensivesurvey Abapical: towards the ápices (of the of Easter Island's sublittoral marine septum) environment. The authors ofthepresent Adapical: opposite to the ápices direc- work had the good fortune of having tion(oftheseptum) access to examples of all the molluscan Aperture, aperturalend: theroundante- material collected during DiSalvo's rioropeningoftheshell. investigation. Other than the few publi- Apex, apical,end: the smaller, narrower, cations that we have been working on closedposteriorendofthetube. (Oliver, 1915, Schilder, 1965, Rehder, Cutting plañe: the plañe individuated 1980,DiSalvoetal., 1988,Raines,2002) by the edge of the shell at the apex no other serious work on molluscs has (excludingseptumandmuero). 50 RAINESAND PlZZINI: Revisión ofthe Caecidae ofEaster Island (Chile) UNITED STA Figure 1. Biogeographical provinces within the tropical eastern Indo-Pacific as proposed by SCHILDER (1965), andas illustratedbyREHDER (1980): Micronesian: M; Hawaiian: H; Fijian: F; Polynesian: P; Rapanuian: R; KermadecIslands: K(addedbytheauthors). Figura 1. Lasprovincias biogeographicas dentro delIndo-Pacífico orientaltropicalpropuestaspor SCHILDER (1965) eilustradasporREHDER (1980):Micronesian: M;Hawaiano: H;Fijian: F;Poline- K sio: P;Rapanuian: R;KermadecIslands: (añadidaporlosautores). Interspace: área between rings, with / Rings, annular sculpture: transverse, withoutmicrosculpture. raised sculpture (equivalent to the Meiobenthic: referred to all interstitial axial sculpture of the normally molluscs living in sediment of coiledgastropods). varyinggranulesize. Septum: closure of the shell at the apex, Microsculpture: usually visible at very asitshedsearlierstages. highmagnificationorunderSEMcan Shell(s): the shell, beached without gas- betransverse,longitudinalorboth. tropod. Muero: small to large prong projecting Spm(s): live collected specimen(s), with fromtheseptum. softpartsand/oroperculum(a). RESULTS SuperfamilyRissooideaGrayJ. E., 1847 Family Caecidae Gray E., 1850 J. Genus Caecum Fleming, 1813 Diagnosis (Bandel, 1996): "The shell of the teleoconch is a small, slightly curved tube orna- mentedonlywithgrowthlines,numerousring-likecollabralliraead/oraxialribs. Theposte- riorendofthetubeisclosedbyaconicalseptum.Theprotoconchistrochospirallyorplanispi- rallycoiled.Uncoilingoftheshellbeginsaftermetamorphosis". Type species (Bandel, 1996): Dentalium imperforatum Kanmaker, 1798 (= trachea Montagu, 1803) fromEurope,MediterraneanSeaandAtlantictosouthernEngland. 51 Iberus, 23 (1), 2005 Figure 2. Caecum rehderi spec. nov. A: holotype LACM 3019, gold coated, length 2.08 mm; B: detailofseptum; C: microsculpture; D: detailofaperture. SEM imagingbyD. Geiger. Figura2. Caecum rehderi spec. nov. A: holotipoLACM3019, metalizado en oro, longitud2,08 mm; B: detalledelsepto; C: microescultura; D: detalledelaabertura. ImágenesalMEBporD. Geiger. Caecum rehderi spec. nov. (Fig. 2) Caecumcf.solitarium;Rehder,1980:31-32,pl.5,fig. 11. Type material: Holotype, LACM 3019; 1 paratype, LACM 3020; 1 paratype, USNM 756269; 1 paratype,MNHN. Materialexamined: 1 specimen,Onetea,Hotuiti(length:2.42mm,USNM756269) (Oct. 1974,leg. H.Rehder);4specimens,PuntaRosalia,eastofAnakena(Apr. 1998,leg.B.Raines), Type locality: In sand collected along thebase ofcliffs at 10-20m, offPunta Rosalia, east of Anakena,EasterIs.,Chile.27°04' 18"S,109°19'45"W. Description: Shell small (holotype withincised annular rings. Aperture cir- measures, length: 2.08 mm; width: 0.42 cular, but slightly constricted. Posterior mm), tube-like, slender, gently arched, end with tapered rim. Septum not semi-translucent to opaque white. Tube retracted, subquadrate lateral outline seemingly smooth almost glassy, sub- inclinedwithelevatededgeslightlyright cylindrical, with posterior end only ofcenterwhenviewedfrontally. Opercu- slightly smaller than anterior end. lumandsoftpartsunknown. Microsculpture nearly obsolete, with Original description ofC. cf. solitar- onlyfineannulargrowthlinessometimes ium Rehder, 1980: ''Diagnosis. Shell mm present under magnification. Anterior small, 2.4 to 2.7 in length, glassy, end somewhatflaredjustaboveaperture grayish-white towhitish, slender, gently 52 RAINES AND PIZZINI: Revisión ofthe Caecidae ofEaster Island (Chile! Figure 3. Caecum cf. solitarium Rehder (1980). A: USNM 756269, uncoated, length 2.42 mm; B: detailofseptum; C: microsculpture; D: detailofaperture. SEM imagingbyD. Geiger. Figura3. Caecumcf.solitariumRehder(1980).A: USNM756269, nometalizado, longitud2,42mm; B:detalledelsepto; C: microescultura;D:detalledelaabertura. ImágenesalMEBporD. Geiger. curved, diameter at posterior end only identified as Caecum cf. solitarium Oliver, slightly smaller than at anterior end, 1915. However, itseems thatRehderun- where the aperture is slightly con- fortunatelyoverlookedseveralkeychar- stricted and somewhat opaque above acteristics within Oliver's description. the aperture; the sculpture consists of The first being the septum of C. solitar- fine, rather crowded, subobscure ium, which Oliver (1915) described as (worn?) annular riblets that gradually "...hemispherical, making an abrupt and slightly increase in strength toward shoulder at the junction of the shell"; the aperture; septum exserted, sub- even the septum of Rehder's specimen quadrate with a slightly convex surface (Fig. 3) could be associated to a tale ty- inclined from an elevated edge at the pology. Oliver also mentions, that the right dorsal sector to the edge of the sculpture ofC. solitarium consists ofsim- posterior rim of the shell at the left ple growth lines, while Rehder refers to ventralsector. the sculpture as consisting of suboscure Range. Kermadec Islands (and (worn?) annular riblets that gradually EasterIsland ?). and slightly increase in strength toward Material. 1 specimen from sta E-27A, the aperture. Other main difference be- USNM756269. tween C. solitarium Oliver, 1915 and C. Measurements (mm). USNM 756269: solitarium sensu Rehder is that the first length,2.42;diameteratanteriorend,0.4". hasanearlyuniformdiameter,whilethe Discussion: Rehder reported a caecid second shows the diameter at posterior from Easter Island, which he tentatively end only slightly smaller than at ante- 53 Iberus, 23 (1), 2005 Figure 4. Caecum solitarium Oliver, 1915. A: Holotype CM M2867, uncoated, length 1.68 mm; B: detailofaperture; C: microsculpture; D: detailofposterior. SEM imagingbyN.Andrews. Figura4. Caecumsolitarium Oliver, 1915.A:Holotipo CMM2867, nometalizado, longitud1,68mm; B:detalledelaabertura; C: microescultura;D:detalletrasero. ImágenesalMEBporN.Andrews. rior end. Furthermore, it appears Re- small rings along the entire tube; and hder did not examine Oliver's type ma- apart from the relativity of the term's terial, because if he would have done, significance, we hold that the difference hewouldhavenoted thatthe two speci- between our specimens and those de- mens have significantly different ante- scribed by the two authors falls within rior ends (Figs. 3D, 4B), and that the the species' range ofvariability, in light, holotype ofC. solitarium isbadlybroken above all else, of the high degree of and lacks the entire posterior end (Fig. adaptation of the local molluscs to the 4). The damage to the holotype is oíd island's distinguishing geo-climatic con- and worn, and possibly occurred in situ ditions. It is known that a number of suggesting thatOliver mayhave chosen species of Caecidae (i.e. C. lightfootae an imperfect specimen as the type and Pizzini, Nofroni and Oliverio, 1994), actually described another in his de- though they have the same general scription. (Scofield, 2002, pers. commu- shape (septum, tube and aperture), nication). Rehder's specimen is, how- could show a very wide range of vari- ever, consistent with C. rehderi, and abilityintermsofthetypeofsculpture. therefore, has been designated as the Remarks: Caecum rehderi seems to be paratype. Although Rehder broke the an unusually fragüe species. Of the five anterior end ofhis specimenwhile mea- known specimens, Rehder chipped the suring it, all the pieces were available aperture of his specimen (USNM for examination. The specimens which 756269)whilemeasuringit,theholotype we found, actually showed a series of has a small longitudinal crack toward 54 RAINESAND PIZZINI: Revisión ofthe Caecidae ofEaster Island (Chile) Figure5. CaecumamydroglyptumRehder, 1980.A: holotypeUSNM757977, uncoated, length 1.67 mm; B: detailofseptum; C: microsculpture; D: detailofaperture. SEM imagingbyD. Geiger. Figura5. CaecumamydroglyptumRehder, 1980.A:holotipo USNM757977, nometalizado, longitud 1,67mm;B:detalledelsepto; C:microescultura;D:detalledelaabertura. ImágenesalMEBporD. Geiger. the aperture end, the otherparatype has the Kermadec Islands, while Rehder a chip in the aperture, and the júnior tentatively indicated the species as author completely crushed another being from Easter Is. In our opinión, specimenwhileexaminingit. according to the present knowledge, C. Geographical distribution: Oliver solitarium is restricted to the Kermadec described Caecum solitarium only from Is. Caecum amydroglyptum Rehder, 1980 (Figs. 5, 6) Caecumamydroglyptum;Rehder,1980:32,pl.5,fig. 12. Typematerial:Holotype,USNM757977;1paratype,USNM757978 Materialexamined:Originaltypes.Holotype,USNM757977;paratype,USNM757978,(Oct.1974, leg.H.Rehder).132specimensinsandcollecWtedalongthebaseofcliffsat10-20m,offPuntaRosalia, eastofAnakena,27°04'18"S,109°19'45" (Apr. 1998,leg.B.Raines). Vouchermaterial:4specimensweredepositedineachofthefollowinginstitutions:LACM;USNM 1018792;MNHN;NHML;NMNZM.273207;NSMTMo73562;AMSC.205278;WAMS13783,and 6specimens(3adults/3juveniles),MPR.13shells,beachofAnakenaBay,onthenortherncoastof EasterIs.,pickedupamongtherockybottomonthewestsideofthebayatlowtide,amongcom- munitiesofDictyotales,withGalaxauraobtusata. (12-1-1995,leg.E.Rolan),MPR. Typelocality:StationE-27A,Onetea,Hotuiti:inpatchofsandabovehightidelevel. 55 Iberus, 23 (1), 2005 Figure 6. Caecum amydroglyptum Rehder, 1980. A: Voucherspecimen LACM, gold coated, length 1.53 mm; B: detail ofseptum; C: microsculpture; D: detail ofaperture. SEM imaging by D. Geiger. Figura6. Caecum amydroglyptumRehder, 1980.A:ElespécimendelvaleLACM, metalizadoenoro, longitud1,53 mm; B: detalledelsepto; C: microescultura;D: detalledelaabertura. ImágenesalMEB porD. Geiger. Original description: "Shell small, the shell being less raised and changing mm from 1.3 to 1.7 in length, curved, their shape, until they resemble very rather evenly cylindrical with the ante- fine growth lines. Microsculpture rior end in fully grown specimens formedbylongitudinalworm-like striae slightly swollen above the aperture; visible at enlargement of at least 180x. glassy grayish white to light orange Septum dome-shaped, slightly raised yellow in color; sculpture consists of overthe cuttingplañe. Aperture consist- rather strong, somewhat distantly ing of a large protuberance crossed by spaced annular ribs that become more slightly raised rings. Operculum and or less obscure in the middle part of the softpartsunknown. shell, with microscopic, longitudinal Remarks: We agree totally with wavy striae that are obscure at the ante- Rehder's conclusions; because we have rior and posterior ends; septum, low, "...been unable to identify this species with dome-shaped. " any published taxon" from either the Additional description: Shell small Indo-Pacific Provinces, the Panamic (mean length: 1.7 mm; width: min. 0.3 Prov. or the Chilean Prov. We have mm, max 0.4 mm), curved, colour foundonlyonespeciesthatresembles C. grayish white. The tube is perfectly amydroglyptum, which is C. vertébrale cylindrical, except near the aperture, Hedley, 1899, from Funafuti Is. It is and its sculpture consists of about 36-40 quite similar to amydroglyptum in terms rings, with some in the middle part of ofthesculpture ofthetube,longitudinal 56 RAINESAND PlZZINI: Revisión ofthe Caecidae ofEaster Island (Chile) Figure 7. Caecum heterochromum spec. nov. A: holotype LACM 3021, gold coated, length 1.42 mm; B: detailofseptum; C: microsculpture; D: detailofaperture. SEM imagingbyD. Geiger. Figura7. Caecumheterochromumspec. nov.A:holotipoLACM3021, metalizadoenoro, longitud1,42 mm;B:detalledelsepto; C:microescultura;D:detalledelaabertura. ImágenesalMEBporD. Geiger. microsculpture and septum, but the tube, a very strong microsculpture Rehder's species shows a much greater consisting of worm-like, longitudinal swelling of the tube above the apertural striae, covering also the top ofthe rings, end, which is crossed by small sculp- while that of vertébrale is an indistinct tured rings, while the adapical part of microsculpturenotsurelycomparableto vertébrale has almost the same diameter arealstriation (pers. observ.). of the tube. In addition, C. amydroglyp- Geographical distribution: This species tum presents, along the entire length of would appeartobelimited toEasterIs. Caecum heterochromum spec. nov. (Figs. 7, 8) Type material: Holotype,LACM3021;6paratypes, LACM3022;6Paratypes,USNM 1018789;6 Paratypes,MNHN; 6Paratypes,NHML; 6Paratypes,NMNZM.273205; 6Paratypes,NSMTMo 73560;6Paratypes,AMSC.205275;6Paratypes,WAMS13780;9Paratypes,MPR. Material examined: 168 specimens: offHanga Nui; and 8 specimens offthe western coastline near Tahai (Dec. 2000, leg. B. Raines). 39 shells,beach ofAnakena Bay, on the northern coast of Easter Is., picked up among the rockybottom on the west side ofthebay at low tide, among communitiesofDictyotales,withGalaxauraobtusata.(12-1-1995,leg.E.Rolan)MPR. Vouchermaterial:39shells,beachofAnakenaBay,onthenortherncoastofEasterIs.,pickedup among the rockybottom on the westside ofthebay atlow tide, among communities ofDicty- otales,withGalaxauraobtusata. (12-1-1995,leg.E. Rolan)MPR. 57 Iberus, 23 (1), 2005 Figure 8. Caecumheterochromumspec. nov.A: paratypefromlotLACM 3022, goldcoated, length 1.53 mm; B: detailofseptum; C: microsculpture; D: detailofaperture. SEMimagingbyD. Geiger. Figura8. Caecumheterochromumspec. nov.A:paratipodelaporciónLACM3022, metalizadoenoro, longitud1,53 mm; B:detalledelsepto; C: microescultura; D: detalledelaabertura. ImágenesalMEB porD. Geiger. Type locality: In sand collected along thebase ofcliffs at20m offHanga Nui, EasterIs.,Chile. 27°07'46"S,109°16'35"W. Derivationoftheñame:Fromthegreekeiepoc,:otherandxpcojxa:colour. Description: Shell small (mean sponding breadth and depth, particu- length: 1.6 mm; mean width: 0.4 mm) larly in the upper portion of the tube with the tube subcylindrical in shape in andneartheaperture,toothersthatlack the abapical part and cylindrical up to anytypeofsculpture;thesetwoextreme the vicinity of the aperture (Fig. 8), representthe limits ofthe species' range where there is a slight swelling, fol- of variability, given that they were lowed by a narrowing of the tube; the found in intermedíate specimens whose apertureisperfectlycircular, simple and rings arebarelyvisible. The microsculp- rimmedby a very slightflaring towards ture also presents a wide range of vari- theoutside. Theseptumisdome-shaped ability, with some specimens not and slightly raised over the cutting showing any trace of microsculpture, plañe. Its sculpture is extremely vari- while the surface of other specimens, at able, ranging from specimens with an enlargement of 30x, presents a approximately50 smallraised rings that microsculpture consisting of a large are separated by interstices of corre- number of very fine, worm-like striae 58

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