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Lips in 1959, the Fltst of Many Field Parties from the University Of

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Lips in 1959, the Fltst of Many Field Parties from the University Of Rev. Siol. Ttq').,41 (3):81 7-842, 1993 A reviewo( the status and biogeography ofthe lizard genera Celestus and Diploglossus (Squamat�: Anguidae), with description of two new species from Costa Rica Jay M. Savageand Karen R.Lips Departmentof Dialog)'. University of Miami, p.o.24 Box 9118. Miami, Florida 33124, USA. (Roo. 17-IX-1992. Acep. 5-ll-19(3) Ah*lId: Two new species of diploglossine lizaros, Celestus hylaiusand Celestus orobius, are described froro Ihe Atlanticlowlands and Cordillemde TaJamanca � Costa Rica, respeclively. The speciesdiffer froro previously Icnown Celestusprimarily in colomlionbut aIso in details � scutellalion (numberoC núddorsal scalerows, lamellaeunder the 4th toe an¡JJCKnumber � preanal scales). The status of Ihe genemCelestus and Diploglossusis reviewed. The alloca­ lion� specieslO eitherof !bese taxa based on tite archltecture � tIteosteoderms is rejected lince Ihepresumed diffe­ rences represent oolOgenelic stages. The tl'l!.dilional cla,ssificauoo whereby species having claw sheatIts are placed in Diploglouusand tItose without in Celestusis resurrected. Underthis amng ement CeJestus is oomposed of 7 MelÚcan and Centl'l!.l American and 16 Antillean fonos whlle Diploglossus includes 6 southem Centl'l!.l and SoutIt American and4 Antillean species. The Hispaniolanendemic genem Sauresia (2species) andWe tmoreNl (1 species)are regarded as derived allies � Diploglossus. Celeslus and Diploglossus are sympatric only in lower Central America but Celestus occurs.in sympatry witIt SallTesia and We tmorena 00 tIte island � Hispaniola. The biogeogmphy oC diplo­ glossines is reviewed wiili special empbasis on tIteCa ribbean region. Two confIictingbiogeogrspbic mode1s (one dis­ persal and onevicariance) tItat purport lOexplain current distribuuonpatte ms (Celestus in Mellico, Centl'l!.l America, Jamaica,Cayman Islands and Hispaniola;D iploglossus inLower Central America,Soulh America,Cuba, PuertoRico, Mootserratand Malpe10 Islands, and ita alliesSallTesÍIA and WetmoNNI on Hispaniola)are examined. Arca cladograms andpredicted pbylogenelic relauonships of circum-Caribbean formsbased on eacb modelare presented. Rejeclion,ac­ ceptllllceor modificauon oC eithermodel must await more detailed pbylogeneuc lIIlalyses. The vicariance model, ho­ wever,il shown lObe tIte most parsimonious explanation oC diploglossinebiogeogmpby based uponavailable data. Key words: Lizaros, Anguidae.Celeslus. Diploglossus, new species, biogeogmpby. In 1959, the flTst of manyfield parties from teras representative of a previously unrecogni­ the University of Southern California set out to zed species. Unfortunately, although considera­ conect herpetological material in Costa Rica ble effort has been expended in searching for 2nd obtained a single diploglossine lizard from other examples of the new form, none have 00- the Cordillera de Talamanca. The specimen en forthcoming and its description is based on was tentativelyidentified as Celestus cyanoch­ the unique type. Ioris Cope,a species thenknown only from !he In addition, sorne two decades ago another Cordillera Central ofCosta Rica, although dif­ apparently undescribed Celestus was discove­ fenog from that forro in details of scutelJation red by Michael J. Corn (MJC). Corn collected and colouration. Since that timemore material five specimens of the putative new form from of C. cyanochloris has accumulated, and the the extensive Río Frio banana plantation in difference ootween it and the Talamancan spe­ Heredia Province. Subsequently, University of cimen remams. This leads us· to regard the lat- Southem California field teams obtained seve- 818 REVISTABIOLOGIA DE 1ROPICAL raI lizards conspecific with Com's specimens made Collowing the procedures defined and/or Crom the La Selva Biological Station oC the illustrated by Peters (1964), with snout-vent Organization Cor Tropical Studies, also in length designated as standard length. Heredia Province, as part oC an intensive study Measurements were made to the nearest 0.01 oC the leaC-litter herpetoCauna (Lieberman mm with dialcalipers. 1986). Colour notes in liCe Cor Celestuscyanochlo­ Although Cor some years Com intended to ris and C. hylaius were taken from colour sli­ describe the Atlantic lowland lizards as a new des projected onto a screen, and diagrammatic species and had collected considerable dataon figures oC colour pattems were sketched from mainland Celestus Cor that purpose, he ultima­ preserved specimens. te1y abandoned the projecL This was due,al le­ Status oC the genera Celestus and ast in pan, to bis uncertainty regarding varia­ Diploglossus on the American mainland: The tion inCelestuscyanochloris, thenknown from evolutionary lineage (subCamily Diplo­ three specimens (counting the holotype),a spe­ glossinae) to which our new species belong cies that he had not seen in liCe, and in part, to containsa single Cossil, Eodiploglossusbore a­ the relationship oC his Atlantic lowland Cono to lis, Crom the Eocene oC Wyoming (Oauthier Celeshu bivittalus Boulenger known Crom a 1982) and 38 extant Conos reCerred to Cour 01' single juvenile from Nicaragua. More impor­ five genera. Diploglossines difCer from other tandy,however, he Cailedto include in bis study anguids in the Collowing combination oC Ceatu­ several scale counts that prove 10 be decisive in res: skinklike in habitus; limbs present but va­ distinguishing among Mexican and Central riously reduced; body covered by cycloid sea­ American members oC the genus, and indeed les that are underlain by osteoderms having a establishthe Atlantic Costa Rican populationas distinctive, peaked, gliding surface (the basal a distinctspecies. non-sculptured surface); a thin, pointed, "quill The present paper serves to describe these pen-shaped" anterior end oC surangular; frontal two new species so that they may be included bones paired; Crontal and maxillary bones in in the senior author's comprehensive handbook contact; and lacking palatine and pterygoid te­ 00 the herpetofauna oC Costa Rica. As pan oC eth and any evidence oC a lateral Cold. Three oC the process oC demonstrating the validity oC the the genera, Ophiodes oC central and southem three CostaRican taxa we haveined exam most South America east oC the Andes, and oC the specimens oC Celeslus available Crom SauresUJ.and Wetmorena oC HiSpaniola are uni­ Mexico and CenualAmerica. We also redescri­ versallyconsidered valid Differences oC opi­ be Celestus cyanochloris based 00 newly ac­ nion exist, however,'concerning the status and quired specimens. Consequendy we provide species composition oC the nominal genera summary characterizations oC all mainland Celestus and Diploglossus; some authors re­ Corms in thegenus anddocument their distribu­ gardthem as Corming a single genus while ot­ tions. In addition we review the characteristics hersmaintain themas distincL and status oC the genera Celeslus and Diploglossus Weigmann,1834 (type by sub­ Diploglossus and develop an original biogeo­ sequent designation oC Fitzinger,1843: 7iliqua graphic hypothesis to explain current pattems fasciatus Oray, 1831) is based on a South of distributiOll. American Cormhaving thefrontal plate in con­ tact anteriody with three shields (a pair oC pre­ frontaIs separated medially by a large frontona­ MATERIAL AND METHODS sal) theand claws on thefingers and toes cove­ red by a scaly (ungual) sheath, so that ooly the Data used in this paper are from specimens tip.is exposed.Celestus Oray, 1839 (monotype: located in several collectioos. Where reference CelestuSstriatus , Oray, 1839) was proposed for is made to a particular example,.the abbrevia­ West Indian members oC this complex having tion denoting the somce Collows Leviton el al. thefrontal contactanteriorly with a single large (1985). shield (the fused prefrontals ando frontonasal). Scalecounts were made witha varietyoC bi­ In addition, as pointed out by Boulenger nocular microscopes under�different magnif1C8- (1885), these lizards have theclaws exposed tions usingasharp teasing needle. Counts were COI'most oC theirlength (i.e. lacka sca1y, ungual SAVAGE el; LlPS: TheC.I 1izud,_ UIII.rand DipIogl088118 819 sheath). Boulengerplaced allof thethen known absent or consists of a single uníl, and one in species of Celestusand Diploglossusin a single which the canal system is extensive and com­ genus for wmch he used the latter name. In his plexo Species having the complex radix were key however, he sepamted theincluded species placed in the genus Diploglossus by these·aut­ into two mlÜorgroups, based on whether or not hors and those in which the radix was simple or they had a large ungual sheath into which the lacking were referred toCelestus. claw could be partially or entirely retracted. The resulting classifícation scheme loo lO a Subsequentwork has added substantially tothe radical realignment of the group, so thats pecies number of species recognized by Boulenger, with exposed claws were placed in each genus. and representatives of his "Diploglossus" are Similarly, each of the putative genera contained now known to occur from Mexico to Brazil and forms with both 000 and three shields borde­ Bolivia, on all islands in the Greater Ailtilles, ring the frontal anteriorly under this arrange­ and on several other Neotropical islands as mento A number of taxa usually regarded as well. During the next 74 years a numberof aut­ closely related by all previous authorS (e . g . hors (Stejneger 1904; Barbour 1910; Burt and cyanochloris and rozellae) were placed in diffe­ Burt 1932) recognized Celestus as distinct, ba­
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