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 reot genera. Because of these variations in the sed on the degree of exposure of the claws, but application of the two generic names we do not Dunn (1939) preferred Boulenger's arrange use binomials in the following several para ment, since the condition of the fronton graphs. asal/prefrontal arrangement did not cOrTelate This interpretation also produced a complex exactIy with the claw differences. After exami and confused biogeographic picture, with ning twenty nominal species from throughout "Celestus" having a distribution in Mexico and the range of the complex, Underwood (1959) northern Central America, Jamaica, the supported Dunn's position. Underwood further Cayman Islands and Hispaniola, and suggested that the ungual sheath had been lost "Diploglossus" disjunctIy occurringin Central in the evolutionary history of this group on se and northwestern South America, eastern vera! occasions and seriously questioned whet Brazil and Bolivia and on the islands of Cuba, her the claw was retractable into the sheath. Hispaniola, Puerto Rico, Montserrat and Most studel1ts of mainland anguids continued Malpelo (off the Pacific coast of Colombia). to recognize Celestus (e.g. Stuart 1963; Peters Fortunately, Strahm and Schwartz provided and Donoso-Barros 1970; Meyer and Wilson evidence that the degree of radix development 1973) although Myers (1973) was inclined to exhibits considerable individual variation. They follow Underwood's interpretation. note that osteoderms from examples of costa Strahm and Schwartz (1977) established tusof Hispaniola "usually" lack a radix, but six that the claws in these lizards are in no way re out of 12 (50%) have a small to moderate one. tractile and presented a new interpretation of In the holotype of atitlanensis a double andre the Celestus-Diploglossus problem based upon duced radix was found on ventral and lateral purported differences in the architecture of the osteoderms but a fully developed one on the canal system of the body osteodenns. As poin dorsal osteoderms. ted out by these authors, the sculptured free Wilson et al. (1986) subsequentIy showOO (distal, posterior) portion of each osteoderm that there is ontogenetic variation in Strahm contains a relatively simple system of canals and Schwartz's sample as well. Individual va thatshows striking variation within,diploglossi riation in radix development is to be expected nes. They calloothis system the "arbor" and in in other species since these authors examined dicated that it is poorly.developed in these li osteoderms from only a single Jizard of each zards as compared to other anguids. A second form in all cases except costatus. The former canal system (the "radix") is usually present in demonstratOO that the nominal species (accor the unsculptured basal (anterior proximal) por ding to Strahm and Schwartz) "Diploglossus" tion of eachosteoderm (that portion overlapped bivittatus Boulenger, 1895 and "Celestus" ati by the free margin of the preceeding scale and tlanensis Smith and Taylor, 1950, are conspeci osteodenn). According to Strahm and Schwartz fíc. The holotype of the former is a juvenile (1977), members.of this complex may be divi whose osteodenns lack a radix and the holoty ded into two groups, 000 in which the radix is pe of the latter is an adult with two kinds of os- 820 REVISTA DE BIOLOOIA TROPICAL teod.erm canal systems (Strahm and Schwartz However, Underwood (1959) and Strahm [1977] examined and illustrated osteoderms and Schwartz (1977) reported that several taken from the two holotypes). Consequently, forros had me claws "half-sheathed". sugges Wllson et al. (1986) concluded tbat the putative tíog lhat interroediates occur. Myers (1973) al- differences in osteoderm canal systems are 00 indicated thataberemay be species differen- subject to ontogenetic variation and have no value in estimating relationships. In light of these variations we concur that tile degree of radix development is lID unreliable diagnostic feature. It should be further noted that no mention is marle ofthis presumed character in subsequent descriptions of new taxa of Celestus and Diploglossus from Hispaniola by Schwartz and rus associates (1979, 1985) and Thomas and A He.dges (1989), wruch are referred to one or the otber genus solely on the basis of phenetic resemblenee to previously deseribed species. Our conclusion, while undermining the Strahm and Sehwartz classiíication, does not resolve whether Celestus is distinct from Diploglossus. In the process oí describing the new species, we became eonvinced that two basic lineages oí diploglossines are represented on the American mainland. These two lineages conform exactly to the traditionaUy diagnosable genera of mainland Celestus and Diploglossus. Mainland Celestus llave exposed daws. a reduced ungllal scale, and a single subungual scale (Hg. lA). Mainland Diplo glossus nave the claws covered for most of their iength by a compressed claw sheath comprised of an elongate and lateral1y expanded ungual scale anó a pair of subungual scales. In addition, there is an enlarged lateroungual scale Iying near me claw base on each side (Pig. lB). Thus, fuere are two scales afOunó tlle base of the claw in mainland Celestus andfive in Diploglossus. Among West Indian diploglossines the same distinctive features are evident (rabIe 1). Claw slleaths are absent in a cluster of eight related spedes from Hispaniola and in a11 Jamaican forms. AH other species from the Antilles, including representatives of the nominal genera Sauresia and Wetmorena. have sheathed claws. The arrangement of ungual, subungual and lateroungual seales is exacUy fue same as in mainland lizards with claw sheaths (Diplo glossus) and without (Celestus). The daw sheath Fig.1. Coodition lOeS diploglossinelizards. A. C1awB featmes appear to be consisten! and invariant for of in exposed (CeleaINa); B. Claw sheathed, more Iban up each species, with those species presumed to be exposed (Diploglos$1IS mllkpwu:úúllS); C. C1aw sbeathed, related (from analysis of other characteristics) on1y up exposed Characler stales andgeographic distributionof recenl diploglossine lizards C1aws Exposed Oaws Sheathed NoOaws 1 Frontonasal 2 Prefrontals & 1 Frontonasal 2 Prefrontals & 1 Frontonasal 1 Frontonasal 1 Frontonasal Celestus Diploglossus Ophiodes CA biviJtatus± CA montanus- CA büoboJus+, u CA/NSA monotropis+, s SSA intermedius- � MICA enneagrammus + CA montisüveslris-, s SAfasciatus± u SSA slriatus- � CA cyanochJoris + SA lessonae+, s MO montisse"ati* +, s SSA vertebralis � CA hylaius+ C delosagra +, s MA millepunctatus+, s SSA yacupoi- p¡. CA orobius+ PRpleei +,s � CA rozellae + � WI strialus - H anelpislus - [ H carraui- H costalus - Sauresia 1 H darlingloni- H darlingtoni- H agasepsoides -, s � H macrolus- H sepsoides-, s � i H marcanoi- H stenurus- &. t¡ H warreni- � J barbouri- � JCI crusculus- � fi J curlissi- J duquesneyi- Wetmorena J fowleri- H haetiana- , s J hewardi- J microblepharis- J occiduus - C = Cuba,CA = Central Arnerica, CI= Cayrnan Islands, H = Hispaniola, J = Jamaica,M = Mexieo, MA = l. Malpelo, MO = l. Montserrat, NSA = Northem South Arnerica, SA = South Arnerica, SSA = Southem South Arnerica, WI = West Indies I.C. = ungual overlap, s = subungualoverlap; naso-rostral contact:+ = present,- = absent e ... = prefrontal and frontonasalfused posteriorly 822 REVISTADE BIOLOGIA1ROPICAL ces in tbe degree of exposure of the claw tip in Diploglossus. on the mainland of tbe Americas. tbose forms having sheatbs. Species purported The several insular species may also beplaced to have the claws half-sheathed are the main unequivocally into one or theother genera, and land forros bivittatus and enneagrammus, the as will be pointed out below, their geographic Antillean microblepharis, and millepunctatus distribution is consistent witb such allocations. of Malpelo Island, Colombia. A review of the These conclusions change tbe generic assign situation indicates that Underwood (1959) ca ments for three HispanioIan taxa (anelpistus, lled two very different conditions half-sheat carraui and warrem) listed in Schwartz and hedo One of these was illustrated for micro - Henderson (1988) from Diploglossus to blepharis (his Fig. 3A) in whieh tbe ungual Celestus. scale is somewhat longer than in the typical It shouId be noted that C. striatus. tbe type Celestus eondition (his Fig. 3C). The second species of tbe genus Celestus. is something of condition is found in millepunctatus (pig. lB) an enigma. The only known specimen was sup where more than the tip of the claw is exposed posedly collected in the West Indies (Gray and the elongate and enlarged ungual scale do 1839; Boulenger 1885). BouIenger, contrary to es not completely cover the more ventral areas Strahm and Schwartz (1977) considered C . of tbe distal portion of the elaw. Two scales su striatus to be related to (not a synonym oí) rround the base of an exposed claw in micro - Celestus occiduus (Shaw, 1802) from Jamaica. blepharis, while five scales surround the Schwartz (1970) and Strahm and Schwartz mostIy eoneealed claw in millepunctatus. (1977) believed that the type must have come Presumably, Strahm and Sehwartz (1977) from the mainland. We are dubious of tbis con thought that bivittatus (and its synonym atitla clusion because C. striatus had three seales nensis)and enneagrammushad elaws similar 10 bordering the frontal anteriorly and 41 seale microblepharis sinee they recorded them as rows around the body. Celestus montanus of half-sheathed. Signifieantly, Wilson et al. Honduras is the on1y mainland Celestus having (1986)make no eomment regarding the presen the deseribed eephalie seuteIlation, and no ce of half-sheathed elaws in bivittatus in tbeir known mainland species has more tban 36 sea redescription. In our examination of available le rows aroundthe body. specimensof tbese forms we can find no exam A summary of generie alloeations of all pIes that differ in sheath development from ot known diploglossine species, those charaeteris her mainland taxa of the Celestus group. We ties involved in determining generic limits in can only conclude that the notion tbat bivitta botb this and previousreviews, and speciesdis tus. enneagrammus and microblepharis have tributions areprovided (TabIe1). In addition 10 !he claw half-sheatbed is based upon misinter tbe eonditions of the claw sheath and prefron pretation or mis-observation, as we would clas tal/frontonasal features we have indieated sify all these speciesas having exposed,unshe whether the nasal smeId is in eontaet with!he athedclaws. rostral (+) or separated from it by a dorsal The condition of the claw sheath in mil/e extension of the first supralabial (-), sinee punctatus however, remains unique and legiti Underwood (1959) and Strahm and Sehwartz mately may be called a partial or incomplete (1977) thought the altemate conditions were sheath. Nevertheless, it has an identieal number intraspecifieally invariate. For those speeies of sheath scales as those mainland Diplo with a elaw sheath we have also reeorded glossus with complete sheaths, and is mueh whether !he unguaI seale overlaps the subun closer in structure lO those specimens having gual (u) or vice versa (s), since this eharaeter the sheathed condition than to any form with appears to show no intraspeeific variation. completely exposed claws. We interpret (his We have not examined the Antillean armngementas indicating its relationship to the Celestus for this feature, but in mainland lower Central American and South American forrns the state is uniforrnIy u. Strahm and stock that ineludes the type species of Sehwartz (1977) observed that microblepha Diploglossus. ris exhibits the s state. In conelusion, we believe tbat the differen The following key will distinguish among ces in claw sheathing pro vide a valid basis for tbe five recognized genera of living diploglos the recognition of two genera, Celestus and sines. SAVAGE & LIPS: The!izard genera Celestus and Diploglossus 823 A KEY TO THEGENERA OF THELIZARD SUBFAMILYDIP LOOLOSSINAE (FAMILY ANGUIDAE) la Four limbs and claws present ...... 2 lb. No forelimbs, hindlimbs reduced 10 flaps, no claws; auricularopening small, hidden Imder 8Ca- les just behindthe angle of the jaw ...... -...... Ophiodes 2a Claws hidden within a scaly sheath (Figs. lB, 1 C) ...... 3 2b. Claws exposed for most of their length; 5 fingers and toes; an auricular opening (Fig. lA) ...... Celestus 3a Four fingers and toes...... � ...... 4 3b. Five fmgers and toes; an auricularopening ...... Diploglossus 4a Anauricular opening ...... S auresia 4b. No auricularopening ...... Wetmo�na DESCRIPTIONS OF COSTA RICAN Honduras in having a single enlarged plateborw SPECIES OF CELESTUS dering the frontal shield anteriorly (threescales Celestus orobius n. sp. bordering the frontal anteriorly); frorn C. en Fig. 2A, 4-5 neagrammus and C. bivittatus of Mexico and Nuclear CentralAmerica in having 21 to 22 la Holotype •• LACM 138540, from the area mellaeunder the4th toeand 66 transverserows near Hortensia, Palma and Fortuna on the of dorsal scales (18 or fewer lamellae, 78 oc Carretera Interamericana, southem slope of the more dorsals); from C. hylaius by having 66 Cordillera de Talamanca, Canton Pérez transverse dorsal and 75 transverse ventral sea Zeledón, Provincia de San José, Costa Rica le rows (76 10 81 dorsal; 84 to 92 ventrals); (1500-2000 m). Collected by R. J. Lavenberg from C. rozellae in having 66 transverse dorsal, and P. H. Starrett, August 15, 1959. and 75 transverse ventral scale rows (73 to 77 dorsals,78 to 86 ventrals). Diagnosis. - This smaIl (standard length 82.85 mm) species appears 10 be most closely Description. - Rostral less than twice as wi allied to Celestus cyanochloris of the de as high, distinctly visible aboye, in contact Cordilleras Central andTilarán of Costa Rica. with nasal; anterior intemasals (= supranasals) narrower than posterior; frontonasals and pre The two forms differ most obviously in colou frontals fused into a single large seute, wider ration, with C. cyanochloris having seattered than long, posterior border straight; frontal na black on the dorsum and that spots flanks do rrow, much longer than wide; frontoparietals 10 not aligo demarcate distinct, light vertical li small, widely separated by frontal; interparietal C. orobius. C. orobius nes as in In addition, has nearly as largeas parietals. separating them wi only 8 preanal seales versus 10 to 12 in C . dely, touching smaller postparietal; parietal se cyanochloris (Table 2) and is a more elongate parated from supraoculars by an intercalated form with the axilla to groin distance 61 % of seale (sensu Myers, 1973); nasal single, nostril the standardlength versusstouter body and axi pierced in posterior part of scale; a suture from lIa 10 groin distance 53 to 58% of standard upper edge of nasal to nostril or missing; a pair length in C. cyanochloris (Table 3). In addition of small postnasals, occasionally fused intoone large postnasal; anterior and posterior loreals 10 colour pattem differences (Figs. 2-3), C . fused; canthal (=superior loreal Tay!or, orobius is distinguished from other mainland of 1956) of similar size, touching a second fiter Celestus as follows (features for the compared nasal, a prefrontal, frrst supraocular, posterior species in parentheses): from C. montanus of TABLE 2 � Scale countslor the threespecies 01 CostaRican Celestus Scales between Supralabials Scales around Middorsal postmental Lamellae Supra- tocenter Preana1s Infra-labials midbody scales and venl of 4thtoe 1abials of eye C. cyanoch/oris AMNH 16290 33 65 73 21-20 10-1 0 7-7 10 8-8 KU 3429 1 34 70 73 25-23 11-11 8-7 10 8-9 KU34292 33 73 76 25 10-1 0 7-7 10 10-9 CRE4590 34 67 75 20 -21 11-11 7-7 10 10-9 � CRE1008 4 32 66 76 23 10- 10 6-8 9-8 UCR7375 32 65 73 24 -23 10-11 7-7 10 9-8 � UCR7376 33 70 77 23-22 10 9-9 O trl Q:I C. hylaius O § FU 31080 30 80 84 24-24 10 -9 7-7 10 8-8 � FU 3108 1 32 80 87 26-26 10-9 7-7 12 8-8 � FU30538 31 80 90 26 10- 10 7-7 12 8-7 FU 75794 30 81 88 27 9-9 7-7 11 8-9 � LACM59202 31 25 9-9 7-7 12 9-8 1:'" KU 125599 33 81 87 27 9-9 7-8 12 7-9 CRE8380 32 80 89 23-24 10- 11 7-8 12 8-8 CRE8381 32 80 86 25-26 9-9 7-7 11 9-9 CRE67 11 32 79 88 25-27 10 -9 8-7 12 9-9 CRE6707 32 76 89 24-24 9-9 7-7 8-8 UCR4544 33 80 92 22-22 10- 11 7-8 12 9-9 C.orobius CRE 277 33 66 75 21- 22 9-9 6-7 .8 7-8 TABLE3 StfJlldard_lUe_ 01 Celestus cyanochloris. C. hylaius. andC. orobius. andtU percent 01stfJll dord14118111 (% SL). standard total groin hesd head head palpebral eat length length % SL axilla % SL forelim %SL body width % SL hindlimbSL % length % SL width % SL height % SL aperature % SL aperature %SL c. cyanchloris ti) � AMNH16290 66.7 142.8 37.4 56 17.3 26 13.2 19 21.2 32 15.7 24 10.2 15 8.2 12 4.15 6.2 2.3 3.5 � CRE10084 80.2 52.4 65 18.5 23 21.8 27 15.6 19 10.7 13 3.4 4 1.6 2 trJ CRE4590 75.85 168.7 222 40.55 54 14.85 20 10.8 14 21.45 28 14.65 19 9.5 13 6.45 9 2.5 3.3 1.3 1.7 p,.. KU34291 juvenile 53.5 135.5 253 30.5 57 7.2 14 6.S 12 9.2 17 11.2 21 6.45 12 4.9 9 2.25 4.2 0.8 1.5 � KU34292 79 156.5 198 46 58 12.8 16 10.1 13 14.1 18 15 19 9.9 13 7.6 10 2.8 3.5 1.2 l.S UCR7375 98.6 237.9 241 56.85 58 14.Z 14 14.3 15 32.85 33 19.7 20 14.05 14 10.05 10 3.55 3.6 1.75 1.8 � UCR7376 98.55 211.9 21S 57.05 58 23.8 24 13.4 14 29.05 30 19.25 20 13.65 14 10.85 11 na 1.55 1.6 ¡:: 81.1 182.1 46.19 57 14.57 18 11 13 21.33 25 15.% 10.71 13 7.97 2.78 3.7 1.32 1.6 ¡;¡ X 226 20 !O eL C. hylaiN.s 1 CRE6707 juvenile 38.5 90.1 234 21.45 56 8.15 21 10.6 28 8.7 23 5.6 IS 3.95 10 2.45 6.4 2.6 CRE6711 74.45 tailblcn. -- 44.3 60 13.85 19 10.4 14 19 26 13.7 18 9.25 12 7.65 10 � CRE8380 106.85 143.7 135 63.8 60 16.3 1.5 15.8 15 27 25 18.05 17 12 11 10.55 10 3.5 3.3 1.45 1.4 CRE8381 91.25 158.7.5 174 SS.l5 60 16.15 18 14.5 16 22.5 25 15.85 17 10.5 12 8.05 9 3.1 3.4 1.4 1.5 8. KU125599 94 168 58 62 11.1 12 11.6 12 14.6 16 15.7 17 9.9 11 8.4 9 2.7 2.9 1.4 l.5 \::! 179 "6' LACM59202 70.5 166.5 236 43.5 62 missing 12.3 17 5.8 8 S- miSllin¡¡ "" UCR4S44 lIS.SS 119.5 140 54.45 64 17 20 11.2 13 25.4 30 15.65 18 10.4 12 9.05 Il 2.9 3.4 1.2 1.4 2" UF30538 105.65 154.3 146 63.6 60 17.55 17 14.3 14 23.55 22 18 17 10.5 !O 8.75 g 2.95 2.8 1.S 1.4 � UF31080 %.15 1.53.15 159 59.3 62 16.75 17 14.7 15 23.3 24 16 17 11 11 8.05 8 2.7 2.8 1.3 1.4 UF31081 103.15 185.6 ISO 60.4 59 18.4 18 16.3 16 26.65 26 17.65 17 12.45 12 9.45 <} 3.25 3.2 1.4 1.4 UF75794 100.25 198.45 198 63.3 63 15.9 16 12.9 13 24.75 25 16.75 17 9.9 10 8.6 9 2.85 2.3 1.35 1.3 X 87.85 139.SZ 178 53.39 61 15.12 17 13.5 14 21.74 2S 15.3 18 9.7S 11 8.25 9 2.93 3.4 1.33 1.5 C.orobUM CRE277 82.85 taH blcn. -- 50.25 61 19.25 23 13.7 17 25.05 30 16.1 19 10.1 ]2 7.25 9 3.4 4.1 l.85 22 e 826 REVISTABIOLOGIA OH TROPICAL A B B. Fi¡.2. Diagntl1UJUltic represernationsof dorsal oolO11rpaUems in Cosla Rican specie. r:i Celutws. A. C.orobiws; C. Itylaiws; C. C. CJQfllJCltloris. e D Fig. 3. Diagrammatic represenlatiOlls of typical donal ooloor pauems in Meloamerican .pecielof Cdestws. A. C. eltlUlagr_(Iateralli¡ht spotI variably present); B. C. rolellae (barring reduced in some large adulta); C. C. morúIlPlWS D. (Iaterallightspots variabl y preaent); C. bjyittatws. SAVAGE &. llPS: The1izard genera Celestus andDiplogl ossus 827 canthal (=fust supraciliary of Taylor,195 6), Cordillera de Talamanca of Costa Rica (1500- and fused loreal; small to medium-sized preo 2000 m) (Figs.4-5). colar; five median supraoculars,2 and 3 borde ring frontal; four lateral supraoculars,anterior most interpreted as canthal as noted by Myers (1973); subocolar nearly 3 times as long as CELESTUSHYLAIUS n. sp. high; a row of four postoculars; nillC suprala Figs.2B,6 bials, six or seven to a point below middle of eye; seven or eight infralabials; mental narro wer than rostral; an azygous postmental; five Holotype. - LACM 138541, from the La pairs of chinshields,fltSt pair in contact with Selva Biological Station, Canton de Sarapiquí, one another and second and third labials; se .Provincia de Heredia,Costa Rica,(40 m). cond and succeeding pairs separated fromeach Collected by James J. Talbot,December 3, other by one to five scales; 66 dorsal scales 1973. from occipital 10 base of tail, 75 ventral scales between postmental and anterior edge of vent, Paratypes. - UCR 4544, KU 125599, UF 30-33 scale rows around midbody; eight scales 75794,UF 30538, UF 31080-81, CRE 8380-81, on anterior edge of vent; pentadactylous; late CRE 6707,CRE 6711, LACM 59202. Locality rally compressed digits with slightly rounded datacan be found in the Materialssection. subdigital lamellae; third and fourth fingers of similar length,with 15-17lamellae; toes relati Diagnosis. - A moderate-sized Celestus vely lQng and slender,fourth longest with 21- (maximum standard length 106.85 mm) with a 22 lamellae,each digit terminating in a claw dorsal pattem of light and dark spots that tend lacking a sheath; upper caudal scales with a to align vertically on the fIank 10 form obscure median keel. bars and paired longitudinal,lateral stripes on a greenish ground colour. It differs most ob viously from the Mexican and Central Colouration in preservative. - Dorsal American species,Celestus enneagrammus and groundcolour olive brown; headuniform olive C. bivittatus. in colouration (Figs.2-3) and in brown; sorne scales dark brown, evenly scatte having 22 10 27, lamellae under the fourth toe, red across neck,trunk and tail; dorsal surfaces versus 14 to 18 in the othertwo species (Thble of limbs uniform brown; definite black-edged, 2). The new form is distinguíshed from the two vertical light lines on flanks; venter uniform otherCosta Rican species,C. cyanochloris and darle grey. C. orobius. by having more transverse dorsal scalerows (76to 81 versus 65 10 73) andin co Measurements in millimeters. - Values in louration.The new form is most similar to C . parentheses indicate measurements expressed rozellae of southern Mexico and northern as percentages of standard length (Table 1). Central America in scalation (Thble 4), but ro - One adult specimen. Standard length 82.85 zellae lacks both the líght-tipped black scales with broken tail; axilla to groin 50.25 ( 61%); scattered over the body,and any hint of dorso forelimb (from axilla) 19.25 (23%); hindlimb lateral stripes; and has very distinct,darle-mar (from groin)25 (30%); head length 10 rear ear gined,vertical light bars on the neck and often opening 1 6 (19%); greatest width of head (an on the flanks. The new species cannot be con gle of jaw) 10 (12%); head height 7.25 (9%); fused with C. Inontanus of Honduras since that diameterof earopening 1.85 (2.2%). form has three plates (2 prefrontals and a fron tonasal) bordering the prefrontal shield ante Etymology. - The name orobius is from the riorly,while all other mainland Celestus, inclu Greek meaning mountain dweller with referen ding C. hylaius. have a single large shield in ce to the montane origin of the only known thatposition. specimen. Description. - Rostral less than twice as wi Distribution. - Known from a single loca de as high,disúnctly visible aboye,in contact lit y in the lower montanerainforest zone of the with nasal; anterior intemasals (= supranasals) 828 REVISTA DE BIOLOGIA 1ROPICAL 102 ,94 90 86 - CELESTUS BIVITTATUS • CELESTUS CYANOCHLORIS [!]CELESTUS ENNEAGRAMMUS .CELESTUS HYLAIUS 12 4CELESTUS MONTANUS @CELESTUS OROBIUS ='A-:1-_+-----¡--¡ 12 * CELESTUS ROZELLAE , 100 /200 300 400 O ! 1 ! t , KILOMETERS 98 94 90 86 82 FJ&. 4. Distributimof mainland specicsof CelutUf. narrower than posterior, fosed in KU 125599; scales occor between postnasals and orbit as in frontonasals and prefrontals fosed into a single UF 31080-81); small to medium-sized preocu large scute, wider than long, posterior border lar; five (rarely 6) median supraoculars, usually somewhat concave to sttaight, (CRE 8380 and 1, 2 and 3 bordering frontal (rarely 2 and 3); CRE 6707 have two small lateral scales split foor or five lateral supraoculars, anteriormost off from this plate in canthal area); frontal interpreted as canthal as noted by Myers narrow, much longer than wide; frontoparietals (1973); subocular nearly 3 times as long as small, widely separated by frontal; interparietal high; a row of four (rarely threeor five) posto nearly as large as parietals, separating them wi culars; nine or 10 (rarely 11) supralabials, se dely, touching much smaller interoccipital; pa ven (rarely six or eight) to a pointbelow midd rietal separated from supraoculars by an inter le of eye; eight or nine (occasionally seven) in calated scale (sensu Myers, 1973); nasal single, fralabials; mental usually of equal width as ros nostril pierced in posterior part of scale; a sutu tral; an azygous postmental; fivepairs of chins re rom f upper edge of nasal to nostril or mis hields, firstpair in contact with one another and sing; a pair of small postnasals, occasionally second and third labials (rarely in contact with fused into one largeposblasal; anteriorloreal of 2; 1, 2 and 3; or 1 and 2); second and succee equal size to posterior; canthal (superior loreal ding pairs separated from each other by one to of Taylor, 19 5 6) of similar size, touching a se five scales. Dorsal scales from occipital to cond intemasal, a prefrontal (or a smalllateral base of tail, 76-81; 84-92 ventral scales b,et scale spUt offfrom prefrontal in CRE 8380 and ween postmental and anterior edge of vent; CRE 6707), fll'St supraocular, posterior canthal 31-33 scale rows aroundbody; usually 12 (ra (fírst supraciliary of Taylor, 1956),and two rely 10 to 11) scales on anterior edge of vent; : loreals (anterior and posteriqr or occasionally pentadactylous; laterally compressed digits posterior and intermediate loreals when four with slightly rounded subdigital lamellae; SAVAGE & UPS: Thelizare! genera Celes/us and Diploglossus 829 86 83 '\---+-----+-----111 101---"\ ---rr-----IIO • CELESTUS CYANOCHLORIS • CELES TUS HYLAIUS @ CELESTUS OROBIUS 9 �----_r------r_----_r------r_--�'__r 9 19 KILOMETERS ______�--�8 L-______L- ��----�-- �------L------� 86 85 84 Fig.5. Distribution of species of Celes/lISin Costa Rica; dotted line indicates 1500 m contour. third and fourth fingers of similar length, dorsolateral coppery brown stripes indicated most often with 17-19 (rarely with 15, 16 or where black scales missing. Dorsum of head 20) lamellae; toes relatively long and slender, coppery brown; tip of snout olive green to light fourth longest with 24-27 (rarely with 22 or coppery brown; upper labial s and suboculars 23) lamellae, each digit terminating in a claw bright yellow-green to yellow-blue; posterior lacking a sheath; aU caudal scales without edges of supralabials and infralabials bordered median keels. by black; thin black line from base of postnasal to comer of eye, along upper edge of subocu Colouration in life. - Dorsal ground colour lars to postoculars, becoming wider and run coppery brown with numerous small black ning along area superior to ear then laterally scales scattered over neck, trunk, and tail, along body. Venter immaculate yellow green; forming black ventrolateral stripes where black colour reaching supralabials, auricular opening colour concentrated from ear to above and sides of body at level of axilla. Iris rusty forelimbs and aJong trunk to groin; pair of brown. 830 REVISTABIOLOGIA OH TROPICAL TABLB4 Range 01 lItllues lor selectedletJtllTU of_inJOnd Cele stus. Means inparenlhau. Species No. Donal No. Ventlal No. Sca1e No. of LameUae ScaleRows ScaleRowI RowsAround on4thtoe Body cyanochloris 65-73 73-77 32-34 20-25 (68) (74.7) (32.9) (22.4) enneagr_ 78-85 81-91 31-36 15-18 (82.2) (86.4) (33.2) (16.5) montllnus 67-72 78-84 33 22-26 (69.5) (81) (33) (24) biyittatus 75-79 76-81 29-31 14-18 (77.7) (79.0) (30.3) (16.8) rozell4e 73-77 78-86 31-33 22-26 (75.2) (82.2) (31.3) (23.5) hyl4ius 76-81 84-92 31-33 22-27 (79.7) (88) (31.6) (24.7) orobius 66 75 33 21-22 (21.5) Colouration in preservative. - Dorsal (12-20%); hindlimb (from groin to tip of lon ground colour olive brown, often with faint gest toe) 14.6-27, x=22.8 (16-30%); head wash of light blue dorsolaterally; a pair of dor length to rear� opening 12.3-18,x=16.0 (17- solatenll light and ventrolateral dark stripes; he 18%); greatest width of head (at angle of jaw) ad uniform olive brown; sorne scales dark 5.8-12.5, x=10.2 (8-12%); head height 8.05- brown, tipped with light blue; evenly scattered 10.55, x=8.72 (8-11 %); diameterof earopening across neck, trunk and tail to give speckled ap 1.2-1.45, x=1.38 (1.3-1.5%). Measurements for pearance; dorsal surfaces of limbs uniform the one juvenile specimen (CRE 6707) can be brown with faint hint of blue-tipped scales; found in Table 3. venter uniform grey, becoming darker grey • • Hylaius blue dorsallyalong flank, upper lip and area su Etymology is from the Greek for perior to earand limbs. forestdweller in allusion to the rainforest habi tatof the species. Measurements in miUimeters. - Values ID Remarks: One of the paratypes (KU parentheses are ranges of measurements ex 125599) of this new form was referred t o pressed as percentages of �tandard length based Celes tus cyanochloris by Strahm and on ten adults. Standard length 70.5-106.85, Schwartz (1977) and their figure of a dorsal x=92.78; total length 119.5-198.5, x=160.8 osteoderm under that name is from this speci (135-236%), seven specimens with broken meno and/or regenerated tails; axilla to groin 43.5- • 63.8,x=56.6 (59-64%); forelimb (from axi Distribution • Atlantic lowland rainforest lIa to tip of longest toe) 11.1-17.55,x=15.9 of CostaRica (40-360 m) (Figs. 4-5). SAVAGE &: UPS: Thelizard genera Celest'"aruI Diploglou", 831 Fi¡.6. Upper,Celest'" hylai",; lower, Celesl'" cyanochloris. 832 REVISTA DE BIOLOGIA 1ROPICAL CELESTUS CYANOCHLORIS COPE anteriormost interpreted as canthal as ooted by Figs. 2C, Myers (1973); subocular nearly three times as long as high; a row of four (occasionally three) Holotype. - AMNH 16270; from Volcán postoculars; lO or 11 supralabials, usually se Irazú,Cartago Province, Costa Rica. ven (sometimes six or eight) lO a point below middle of eye; nine (rarely ten sometimes Diagnosis. - A reIatively small lizard (lO eight) infralabials; mental a little wider, but of 98.6 mm in standardlength) most similar lO C . less area than rostral; an azygous postmental; orobius of the Cordillera de Talamanca of five pairs of chinshields, ftc st pair in contact Costa Rica,but differs in having scattereddark with one aoother and second and third labials; spotting, and 10 lO 12 preanal scales versus de second and succeeding pairs separated from fm ite,black-margined, vertical light bars on the each other by one lO five scales; 65-73 dorsal flanks and eight preanal scales in C. orobius. scales fromoccipital lO base of taíl, 73-77 ven Celestus cyanochloris may be separated from tral scales between postmental and anterior ed all other mainland Celestus as follows (charac ge of vent, 32 lO 34 scalerows around body; 10 ters for the compared forms in parentheses): scales on anterior edge of vent; pentadactylous; from C. montanusof Honduras in having a sin Iaterally compressed digits with slightly roun gle largePIate bordering the frontalshield ante ded subdigital lamellae; thirdand fourth fm gees riorly (three plates); from C. bivittatus of of similar length, both with 16-18 lamellae; to Nuclear Central America and C. enneagram es relatively long and slender, fourth longest musof Medco in having 20 lO 25 lamellaeun with 20-25 lamellae, each digit ending in a dee the4th toe (18 or less) and dorsal scales in claw withoutsheath; upper caudal scales with a 65 lO 73 ttansverserows (75 or more); from C . median keel. rozellae of northern Central America and C . hylaius fromCosta Rica in the numberof trans Colouration in lite. - Dorsal ground colour verseventral scalerows, 73 lO 77 (78 or more). tan anteriorly becoming darker posteriorly with Differences in colouration (Figs. 2-3) provide rusty brown tail; head light brown with few additional means to distinguish this species scattered black spots; tip of snout pale yellow fromits congeners. tan; black scales scattered irregularly over body, often forming short irregular stripes Description. - Rostral less than twice as wi across dorsum; not concentrated as lateral stri de as high, distinctly visible above, in contact pes running length of body; some blackscales with nasal; anterior internasals (= supranasals) tipped with lighter tan; limbs appear mottled narrower than posterior; fronlOnasals and pre with brown, black and light tan, one specimen frontals fused into single large scute, wider with leading edge of forelimb yellowish-green; than long, posterior border somewhat cOllcave; supralabials,infralabials and ventrolateralareas frontal narrow, much longer than wide; fronto of neck below level of ear ivory lO white, one parietals small,widely separatedby frontal; in specimen with considerable black flecking in terparietal nearly as large as parietals, separa white colouration fromear posterior lO axilla; ting them widely, lOuching much smallee inte posterior edges of supra- and infralabials bor roccipital; parietal separated from supraoculars dered with black; thin black line from comer of by an intercalated scale; nasal single, nostril eye along dorsal edge of suboculars; ven ter pierced in posteriorpart of scale; a suturefrom greenish-yellow. upper edge of nasal lO oostril; a pair of small postnasats; anterior loreal slightly smaller or of Colouration in preservative. - Olive brown equal sizelO posterior; cantbal (=superiorloreal dorsally; sides of head, neck and trunk fadiog of Taylor, 1956) of similar size, lOuching a se lO light brown and becoming blue-grey venteo cond internasal, prefrontal, first supraocular, laterally; two specimens (KU 3429 1, CRE posterior canthal (=first supraciliary of Taylor, 4590) with typical colouration but with nume 1956), and two loreals; small lO medium-sized rous black punctations evenly scattered across preocular; five median supraoculars, second brown areas of trunk and tail, and two faint and thirdbordering frontal, first excluded from dorsolateral stripes weakly indicated where contact; four (rarely three) lateral supraoculars, black spots absent, one specimen (CRE 4590) SAVAOE &. LIPS: TIlelizard genera Celestua and Diploglossua 833 has bluish-green dorsal ground colour, beco Cope(18 94)described the calouration ofC. ming yellow-green posteriorly; venter uniform cyanochloris in preservation as follows: "co grey (blue-grey in CRE 4590). lour above light golden-green, with several in distinctlongitudinal rowsof paler spots as large Measurements in millimeters. - Values in as a scale, mingled with as many brown spots parentheses are ranges of measurements ex as large as a scale. Lower surfaces blue, paler pressed as percentage of standard length from on cmn and tail".Preserved specimens from the mx adult specimens (AMNH 16290,CRE 4590, Cordilleras de Tilarán and Central agree well CRE 10084, KU 34292, VCR 7375-6). with this description although usually having Standard length (snout-vent) 66.7-98.6,x= 83.2; more olive than green dorsally. One example total length 142.8-238, x=183.6 (198-241 %); (CRE 4590), as described in the section on co louration aoove, matches closely the colour of axillalO groin 37.4-57, x= 48.4 (54-65%); fore the holotype, whose calourhas probably dulled lirob (from axilla) 12.8-23.8, x= 16.9 (14-26%); somewhat over the last97 years in preservati hind timb (from groin) 14-33; x=23.4 (17- ve. AlIavailable examples from mesemountain 33%); head length lO real' ear opening 15-19.7, ranges have a definite bluish casto In preservati x=16.7 (19-24%); greatest width of head ve C. hylaius is olive-brown above and charac (behindcomers of mouth) 9.5-14, x=H.33 (13- teristically marleed with scattered darle scales 15%); head height 6.5-10.9, x=8.6 (9-12%); tipped with biue. Celestus orobius has distinct diameterof ear opening 1.2-2.3, x= 1.62 (1.5- vertical lateral light bars. 13.5%). Measurements for the one juvenile It should be noled thatin JiCe C. cyanochlo ris has a to brown dorsum marleed with (KV3429 1) canbe found in Table 3. tan scattered blacle scales and the venter is gree nish-yellow. In contrastC. hylaius a bronze Etymology. a Thename is from the Greek has to copper dorsal ground colour with a defmite cyan (darle biue) and chloris (green) with refe rence lO the holotype's calour in preservative. greenish casto numeroos bicoloured (blacle and greenish) scales, and a greenish-yellow ventero In preservative theoverall hue of C. cyanoch Remarks. - This species was frrs t described by Cope (1894) along with 11 otiler new Costa loris is bluish-green wmle C. hylaius is brown above and blue-graybelow. Rican forros based on material in the Museo Nacional de Costa Rica loaned to him by The type locality and colouratioo effectively George K. Cherrie, who at thal timewas a taxi rule out the possibility that Cope's montane example was a representative of the lowland dermist and conector for me Museo. The UJli... que typesof two of these species (Hyla · c�rre i C. hylaius. It also seems highly unlikely that and Hyla chrysops) cannot be located in'any the Volcán Itazó specimen is conspecific with the Cordillera de Talamanca species (C. oro caUectioo andmust be regarded as lost, but all bius) because of differences in colouration. of me others reached the American Museum of the Natural History by some circuitous route and Consequently, we follow TayIor in using the name C. cyanochloris for theCordillera Central remainhoused safely there. Taylor (1956) was thefirst worleer to refer lizards and conspecifics from the Cordillera additional specimens to C. cyanochJoris. His Tilarán. material (KU 34291-92) is from the slopes of VolcánIrazú in the same mountain range as the holotype (the Cordillera Central). Disiribution. - Premontane zone of the Cordilleras de Titarán and Central of Costa Taylor thought that theholotype ofC. cya nochWrishad bren lost OOt assumed that bis Rica (1200-1550m) (Figs.4-5). material was conspecific with it Cope's ori It soould be notedthat thespecimen, now lost, reported as Celestus steindachnerii by ginal description did not provide data that ' would unequivocally distinguish the holoty Cope (1893) from Boruca. Puntarenas Pro vince, Costa Rica (550 m) remains an pe from other Costa Ricanforros but OUT re enigmá examination of the holótype confirms as pointed out by Taytor (1956). Cope deseri Taylor'susage. bed it as having two prefrontals, 36 rows of 834 REVISTA DE BIOLOGIA1ROPICAL sca1es around tite midbody, and thebody scales C. cyanochloris: CostaRica: Cartago: Vo lcán smooth anteriorly but stricted and lacking a Irazú (AMNH 16290, holotype); Heredia: Isla median keel posteriorly and on the taH. Tbe Bonita, Vo lcán Poás (KU 34291-92); rus t listed Ceature (Table 4) eliminates all main Puntarenas: Monteverde (CRE 4590, CRE land Central American species, except 10084,UCR 7375 -76). Celestus montanus (Honduras) fromconsidera tion. lbe second eliminates C. montanus and C. enneagrammus: Mexico: "Tehua!ltepec" all known Costa Rican Corms as possible cons (USNM· 30189); Vera Cruz: Orizaba (USNM pecifics as well. In addition C. cyanochloris 6342, USNM 6p(3), 5 km SE Perote (KU and C. orobiushave well-developed median ke 26733); Jalapa ( MCZ 2848); Oaxaca: Sierra els on the caudal scales. If the p�ence oC two Mixe, 0.8 km W (by road) Totontepec prefrontals is regarded as a possible anomoly (UTACV 12215- 19, UTACV 14596); 3.6 mi W the scale count and tite structure oC the poste (by road) Totontepec (UTACV 7727-29, rior dorsal and caudal scales are similar lo tho UTACV 8388, UTACV 9520), 4.9 km W (by se oC Celestus enneagrammus (Mexico). Even road) Tolontepec (UTACV 10278). iC the localitydata Cor Cope'sexample are erro neous identity with C. enneagrammus seems a C. hylaius: Costa Rica: Alajuela: Río Cuarto remote possibility. ConsequentIy, we conclude (UCR 4544); 1.6 km W La Fortuna (KU in agreement with Taylor that an undescribed 125599); Heredia: La Selva (CRE 8380, ho species oC Celestus may await rediscover from lotype; CRE 8381; CRE 6707; CRE 67 11); Río southwestem Costa Rica. Frío (UF 75794, UF 30538, LACM 59202); 3 km SE Puerto Viejo (UF31 080-81). C. montanus: Honduras: Cortés: "few km S MAINLAND SPECIMENS SantaElena" (LSUMZ36659, UTACV 9443). OF CELESIUSEXAMINED C. orobius: Costa Rica: San José: Intera merican Hwy. between Hortensia and Fortuna C. bivittatus: �I Salvador: Ahuachapán, Finca (CRE 277, holotype). El Imposible (KU 18048); Guatemala: Atitlán; probably San Lucas Atitlán (MNHP 5206); C. rozellae: Mexico: Chiapas: Palenque Nicaragua: Matagalpa: Jérico (BMNH (*USNM 113526); Vera Cruz: Jesús Carranza 1946.8.29.37); Chinandega: Volcán Chongo, (KU 27514-1 5); Guatemala: El Petén: Tikal Cordillera de los Marrabios (KU 194658); (UF 13726); 20 km NNW Chinajá (KU 55861- Honduras: Intibucá: La Esperanza (KU 62). t 194665-68, KU 194669-78); Lempira: * Gualcince (KU 194679); Yoro: Montafia de MJC notes Macuzul,aboye El Portillo (FMNH236386). t Wilson et al. (1986) data A KEY ro THE MAINLAND SPECIES OF THE GENUS CELESTUS la. . Frontal shield bordered anteriorly by a single large plate (fused prefrontals and fronloparie- tal) ...... 2 lb. Frontal shield bordered anteriorly by three plates (two prefrontals and a fronlonasal); small dark spots on dorsum and flanks, sometimes with large, dark-outlined, lime-green light spots on flanks...... Celestus montanus (Honduras; 915-1373 m) 2a. 14-18 lamellaeunder 4thtoe ...... 3 2b. 20-27 lamellae under4th toe...... 4 SAVAGB & J..lPS:TIíi:lizaJd generaCeleifu lDdDiploglou llll 835 ...... 31. A pairo f distinctdorsolateral light stripes �...... '...... ; •.... • ...... f.;.; ...... Ce lestw biwnatus (Guatemala10 Nicaragua; 1roO-1882 m) 3b. Dorsum light brown, sharply contrasting with darker·fJanks'that may eitheruniform be 'or mar- ...... ked with laIgelight spots ...... � ... ; ...... Celestwenneagr ammus (Mexieo; 1200-1900 m) 41. 78 or more tFansverse TOWS of ventral scales; 73 10 81 transverse rows of dorsal scáles: caüdal ges lack a distinct median keel...... 5 4b. 77 orfewerbnsverse tows ofventral scales; 65 10 73 transverse rows of dorsal scáles; caudal . scáleswith a: distinetmedian keel ...... � ...... óo ...... :.· ...... 6 - 51. Dorsal and lateral pattem includes scattered, light tipped blac� scáles tbat sometimes fórm mort longif.Jadinal (X" verticál lines; side of neck witÍlOut vertical dark and light bars; á'púr of longitudinalgreen im dorsolateral and black ventrolateral stripes present...... " ...... ;; ...... Cele stw hykn'Us(Costa Rica;40-36( 111) , Sb. Dorsumuniform or With some black ftecking; distinct altemating light and dark bars on side'of . neckand usuallyon t1anks; no indication of longitudinal stripes: ...... ; ...... Celestw rozelloe (southem Mexico. Guatemala, Belize: 2-480 m) 61. 10-12 preanal scáles; body pattem of scatteredblack scáles .. :�: ...... ó ...... : ...... ; ...... Cel estw cyanochloris (CostaRica; 1200-1500an) 6b. 8 preanal séaIes; body of scattered bIack scáles'dorsallyand definite black-tnaIgine(t light:bars on flanks...... Celestw orobius (CostaRica;' 15 00-2000 m) DISTRIBUTION ANORELATlONSIDPS Celestwbiwtt atus and enMagram mus show si milarities in scale featlD"es but are·strikingl y The species of Central American Celestus different in eolouration, whereas C. cyanoch� are completely·állopatric 10 oneanother in dis loris and C. orobiu s appear to be allied. tributiono'l'Wo species (C. hylaiusand C. roze Celestus montanus does not appear 10 be cl� llae)are found only in hliínid lowland fores ts sely related 10 any o�� form álthough it re . of the Atlantic slope below 500 m in áltitudC. sembles the upland Costa Rican forms in some The$C lizards do not párticularly resemble one' scále features � . another and their ranges are separated by a dis . . A cladistic analysis of the seven Central tance of some 500 km . The remaining five American species here placed in Celestus was forins are frorn the .uplands above the 1200 m undertaken using PAUP (Swofford, J98S) ha contolD" and 1lJ'e' distributed on isolated upland sed upon eight characteristics (scale rows masses,in a IUtear fashion from north 10 south around body, number of dorsal scale rows, as follows: Atlantic drainage Slopes eastem and numbee of ventral.scáles, numbee of preanals, southeastem Mexieo (enneqgrammus), modera naso-rostral contact, number of fourth toe la te elevations C American meUae, color of Ibe' Nuclear. entral condition of frontonasal. and pat highlands from Guatemala 10 central Nicaragua tem). Monophyly was based opon the unSheat (biwttatusl. montane Atlantic Slope Honduras hed condition of the claws whicll Underwood (montanus), Cordillera de TIl6ran and Centrál (1959), Strahmand Schwartz (1977)and w e re of Costa Rica (cyanochloris) and CordiUera de gard as derived from the sheathed condition in Talamanca of Costa Rica (orobius). This se Diploglossus. Diploglossus bilobatw and D . quence doe,s not appear to·reflect an easüyde monotroP4 we� used as outgroups. These we termined set of phylogenetic relationships. re especial" appropriate choiees because the RBVISTA DB BIOLOOIA fo1!'m-� has a single shield. in me preoonw afea 3. "Celestus" arose from me ancestorin Central America, probably in association witb (3large frontoM.sa1) ane .dle latt.er ihree (2 pre.- fn:'I!1I�� ,� üoníon.M;fll), Inooilicases the Miocene or eMlier "�,ing!> mud clcsings" aMlysis produced M I.mre80lved . polychotomy oíme PMamanian ·Portal of aU seven species. strongiy indicating diat ro me omerkinds oí data (e.g, molecular) s:re neC\ 4. "Diploglossus"retained me ancestral cbarac aed f.O estabH51n a robust Ilypothesis oí rela� ter states and remained in SOllth America, tionships fOl' mainland Celestus, althougbnow ranging nOM 10 GuatemaJa . 5. "Diploglossus" invooed the Antilles by over water tnIDsport or island h opping via the BIOOEOORAPHIC CONSIDERATIONS LesserAm illes.lO theGreater Antillesexclu siveof Jamaicaand !:heCayman Islands Sll"abm and Schwartz (1977) proposed a googmpbic model fo! dle major featm'es in !he 6. "CelesKus" subsequently dispersed to Mstoty ol diploglo$<�ioo evolution based on the Jamaica ami the Cayman Islands via ove!' c,,"IDcept . that the ancestor of the living genera water transponand menee10 Hispaniola wa� oí South American stílCk having sheathed c1aws, ana M amicuw opening. Although ot� ,. Sauresia and Wetmorena evolved from her fea tures were attributed ro me presumed an� Celestus in. situ00 Hispaniola, and seconda· cestor (radices. ·3 shields boroering me fronw rily regained elaw sheatbs. anterlody and a n¡;¡so..rostta! oontact) all of the� se conditions exhibit intraspeci,tic variation in This scenano is a classic disperal explana one 01' more contemporary species and nave tion of a distributional pal:tem involving a cen questionable utility in establishing relations ter of origin (South America), a migration (to hips. h shoold be notro oowever. tbat a majo Centml America) and two episodes oC overwa rity oí �ie$JUlv e a single lm'ge frontonasal ter dispersion to the Westmrues (pig.1). It al' shield and mootfoons nave the nasal separated pears to support the 10fig-held vrew (Simpson ftol'll tbe ros!laJ Table 1). 1942, 1946; Darlington 1951; Williams 1989) Tbe Sttahm and Schwam biogeographic that me biota of the Orearer Antilles arose by mooel is briefly smnmarized oolow. The reader waif. dispersa! · from the mainland and subse shol.'lld bear in mind oowever. thataccording lO quent island hopping along me Lesser Antilles out a.rW)'sis ilieit concepts oC "Celestus" and are. However, before either the Strahm and "Diploglossus" are unnaturnl (non-monophyle Schwartz scheme or its congruence (Oi' lack tic) tau, smce Strahm and Sc:hwartz placed ID9 thereof) with me general dispersal hypothesis dhiduals ohhe :!laMe$pecies mto different ge� can 00 evaluated, itis necessary to eluddate di nera andmal both nominal genera contain spe ploglossine distrlbution as required by our re cíes witb and wimout daw sbeatbs. Their emp� evru.uation of me species rontent of Celestus hasis on me apparently tmreliable ano variable 8..'fldDiplog lossus. osreodermal ca."ull systems as an indicator ef As delimited bere, Diploglossus occurs in relatioMhips is principaUy re.spon sible · for this Soum America, Lower Central America, and situatien, as they define!! "Diploglossus" as ha� on me islands oC Malpelo. Montserrat. Puerto viog radices and "'Celestus" as lacking mem. Rico and Cuba. Sttahm and Schwartz's inelu SimUady. Sauresi.a and We mwreM were·rela- sion oí Upper Central Amenca in the range of lO "Celeslus" on mis Oasis alone. The Sil me gemAS is .based opon reference of Celestus lient featul"eS of thekmodel are: atidanensis·(a junior synooym oí Celestus bi vittatus) lO Dipioglossus. Simil.arly, Hispaniola 1. A widespread tropical American ancestor oí was ineluded in tile disttíbution oí Diplo Soum American origin differentiated into glossus based upon me p1acement of Celestus ilie modero genera wammi in me genus (tile allied Hispaniolan species. C. anelpistus and C. carraui.Wete des 2, Ophi.t:>dtes evolved in situ in South Amerlca cribeds ubsequent to me 1977 paper). 837 . 100. 90 80· 70· 60· �er study of tIle relationsbips of tIle Wd · 30 eoo Indianforma. The modifications in generic eharacteriza tions and composition oútlined above do not ·· substantielly modify the overallmodel proposed by Strahm and Schwartz. A revised dispersa! model bised on our systematie reallocations involves a migration' from South to Central America and two . overwater dispersais 10 the We st Indies; Gne along the Lesser Antillean ehain aqd.. the' other frQm Central America 19 Jamaipa and then lO HispaJÚola. This conforma to � esse.ntial features of tIle Strahm and Schiartz hypothesis, and is presented in the 80· 70· 60· form of a eladogram oí .,ea relationships (F1 8. 9). This hypothesis lequires a mínimum of one 30· 400 800 vicariant and seven dispersal events, thrce of which are major.It could be corroborateél if the pbylogeny ' of diplogiossines were determined to becoogruent wi th the areacladogram, since such a lypology is predicted by their modelo Unfórtunately such a test of the dispersal model must await a rigorous cladistic analysis of diplogiossine phylogeny (see below) whio1l ií beyond the scope <#tIlis papt. ,w. donote with mote tJ,uma. caSual .. in�i. ··tbJt tbe distribution of diplog10ssines conform.. clQsel)' to that 'expected from Guyer and Savage�. (1987) vicariance model of Caribbean 1?iogeop'aphy (Fig. 10)based upon an interpretation fi¡. 7. DUpenalhypoIheaiJ d etiploatomne biOSeo¡rapby of recent for Muo.merica .rut die We at Inetica. UP, " • CELESTUS • * DIPLOGLOSSUS _ SAURESIA/WETMORENA � OPHIODES 800 1600- I ! HllOMETlAS Fig. 8. Distributions oí extant genera oí diploglossines. Ctlestus is sympatric wilh Sawuia and We tmo7l!M overmuch oC Hispaniola. 4. The fragments that now form the 6. Reconnection of Nortb and South America southwestern portion of Hispaniola and by the uplift of the Panamanian Isthmus Jamaica remained associated with Central resulted in sympatry between formerly Americafor tbe longest time disjunct, allied taxa via dispersa!. 5. Hispaniola is a composite istand formed by The present distribution of diploglossines accretion of several parts, with the and their apparent relationships show a close southwestem portion added most recently correlatianwith this series of events as follows: SAVAGB & LIPS:TbB 1izud paera Celu,,,. andDiplo,1o#u 839 ______.1 _ _ ___ lIGA " .. CM e .. • -lA - .. -- � ----- LA e H .. ""¡le / NCA_J � oJ.CA� � " " " NCA."II H/C.CM'.. 'IA '�LA .... • H ··1 C NCA'PGAIIA by bioaeoarapbic hypothcse a. Pis. 9. CacJosnmalreJátiOlllbipl al Celu"" and Dipló,lou". predi<:ted Left, diJpenal model;vicariance riJht, modeLD = majardiJpeml evClll, d = minOl'diJpeml evClll, V '" vicariancc event; anowl indicate diapenUI,bus vicariancceveotI; e = Cuba, CH = cenltalHillplDiola, H = HüpanioJ.a. l = lamaiéa.LA = Lea.erAmilles. M = Monbernt.NCA '" NuclearCenlral AmeriC!l. PR = PuertoRico, SA = South AmericI.SH = lOUlhem HiIpanio1a. See texl fOl' dilcullico. 1. A widespread Ucestot resembling Diplo 6. Diploglossus and Celestus now overlap in glossus in haviog a claw sheath ranged over Lower Central America following, the uplift tropical America of the Panamanian Isthmus and their dispersal (north or north andsouth) inlOthe 2. Fragmentation ofthe proto-Greater Antilles region. split this stock intwo, a southem component in South Americá and on the fragments that This model requires only three dispersal becamePuerto Rico, �ntral HispaniQla and events lO produce the cUlTent pattem (Fig. 10). Cuba plus a noithem unit comprised of It is also corrooorated by the aistribution of � . Central America, future southwest" ern other groups of Antillean species, including Hispaniola andJ�ca anoline lizards (Guyer and Savage 1987), butterflies (Miller . and Miller 1989) and 3.·�Celestus differentiateC:l in the northem atea; Eleutherodactylus (Hedges 1989), suggesting speciation within Diploglossus resulted in that the pattém is a general one and that the . me origin ·of .the Antillean taxa Sauresia and modelmay have general·explanatory power. We tmorena on the isolated fragments that Selection of one of the competing hypothe fotID central Hispaniola . were 10 PuertoRico, ses of diploglossine biogeography over the andCuba otber (or rejection in favor of some other 4. The fragments that were to become hypothesis) is dependent upon a cladistic ." . so�thwestern Hispaniola. and Jamaica analysis of diploglossine phylogeny basedo n n osteological andlor molecular charaeteristics . . �ly split,off trom Ce tral America, Rejection or acceptance of the dispersal or Q3t1')'ing Celestus·with th� . vicarlance · hypothesis is depéndent upon how 5. Accré,tion to the southweStem portion ·of closely the phylogram conforms to the , HispáriiolaprodUced the pí'esellt sympalf)' of cladogram of atea relationships requiJed by the Celestus withSauresia_ and We tmorena competing hypotheses (Fig. 9). 840 RBVISTA DB BIOLOGIA TROPICAL PAL.OC.N. EOC.N. ACKNOWLEDGEMENTS We are indebted 10 !he following curators for their cooperation in providing specimens �d data for inclusion in this study (abbrevia nons for catalogued specimens in parentheses): D. L. Auth.University of Aorida (UF);J. A. Campbell. Univetsity of Texas at Arlington OLIGOC ••• .ARLY Mloe.N. (UTACV); W. E. Duellman, University of Kansas (KY); R. W. McDiarmid, National Museum of Natural History (USNM); C. W. Myers. American Museum of Natural History (AMNH); D. C. Robinson. Universidad de Costa Rica (UCR);: D. Rossman, Louisiana StateUniversity (LSUMZ); H.K. . Voris, Field PUOC ••• Museum of Natural History (FMNH); J.W. Wright, Museum of Natural History of Los �geles County (LACM). Special thanks go 10 MichaelJ. Com of Lake College,Forest whose notes and counts and measurements of additional specimens were most useful in Fil. 10. Ouyer and Savale'. 1981 vicariance modol of evaluating the status of our specimens. We �ri�,*!.eograph� al an expl�tion d diploglossine o appreciate the aid of Maureen A. Donnelly, dlUributíOD m the· rellon Lipt lupple distribution of . American Museum of Natural History, who DlplogIDl,,,,·like aneellOr (paleocene) and Diplo,IDIS"" provided supplementary data needed for darle1Iipple , distribution al e,I",.". C= Cuba, H = cen� Hispaniola,J • Jamaica, POA = Proto-Oreater Antilles PR outgroup·comparisons, of Michael Fogdeo who = Puerto Rico, SH = lOIIthem Hispaniola. Scale al �tral kindly donated color slides of C. eyanoehloris American IIlhmuI in Paleoc:ene and Proto-Oreater Anti1le1 and C. hylaius. and of Joseph B. Slowinski for exallerated for purpolel of illultration. Prelent constructing the parsimony cladogram. We diltribu�OD � and Diploglo"", and itl allies . e,I",,,, detailedm. FlII.1-8. Seo teXt forfudher explanation. thank M. A. Donnelly ánd C. Guye.r for comments on the manuscripL In the �nterim. we invoke the principie of RESUMEN biogeographic parsimony as our basis for favoringone of thesehypothes es over the other. Sé describe dos especiesnuevas de l8gartijas TIlis. principIe.iStates that.when two competing diploglosinas. Celeitus hylaiús y Celestus biogeographic hypotheses are compared. the one orobius; d� las bajuras del AtÍántico y de la conforming most closely to geographic and Cordillera de Talamanca. Costa Rica. geoIogic data and requiring the fewest ad Me respectivame AIll�as difieren de las dispersal events should be favored (Le. it ls most nte. descritas previamente el género CelestúS parsimonious). In this case there are no major para fundamentalmente en la coloración, como differences in the geologic basis or .geographic así en detalles de las escamas (número de filas relationships iQvolved in thetwo hypotheses. medio-dorsales. lamelas bajoel cuartodedo y/o although the dispersal model implies a greater número de escamas preanales). Se revisa el recency of events (e.g. after fragmentation of and estado taxonómico de ambos géneros. Se movement by the Greater and Lesser AntilIes rechaza la ubicación de especies en alguno into more or less. their present positions) than de esos taXones que se basa en la 81quiteclura de . does thevicarianc.e one. However. the vicariance los oSteodermos debido a que las 'supuestas hypothesis provides the. more parsimonious diferencias demOstraron ser en realidad explanation of diploglossine,biogeography since, v�ación ontogén\ca. cambio. se &\.nueva it r"9uires on1y tlu'ee dispersaI events compared En validez a .Ia clasificación tradicional q.ue to elght Strahm Schwartz dispersal for the and· ubicaba en Diploglossus especies hypothesis (Fig. 9). las con vaina SAVAGB " UPS: The lizarcJ pIlera C,'ulU8 andDiplOf lOllU8 841 1982. en las uftas'Y Iascatentesde vaina en Celestus. Gauthier, J. A. Fossil lUmOllllrid and an¡uid lialdl from &he Boceae Wa .. .ou&h".l AsC, CeleslUSt tiene 'Siete especléS en Mé Xico y eady tCh FormatlClll, Wyomin,, and ' a revision el the An,lÍioidea. América ' Central , 16. en las Antillas. y Conlribulion. 10 Geolo,y, Univerlity of Wy omilla Diplogolssustiene seis en Centroy SurAmérica 21:7-54. y cuatro 'fIl las Antillas. Los géneros endémicos J.B. de Isla Bspaflola, . Sau.resia (dos esp�cies) y Oray, 1831. DlIICIipdanof a De\\' ¡enu.al apJduurean animal, discovelld by &he Jame. •• We tmorena (una) son parientes derivados de late Hunter. BICI In New HollancL Zooloaica1Mi.celIan y 1:14. Diploglossus. CeleslUS ,y Diploglossus solo son simp4tricos en América Central, pero Celestus Gray, J. B. 1839. A Qalalo,ue of &he .lender-ton,ued coincide con Sauresia y We tmorena en la .auriaRl, wi&h delCriptionl of many new lenera and Ipecie¡; Annall and Ma,am. 01 Nalural Hinory Espafiola. Se revisa la biogeografCa de los . diploglosin()s hacienc.tp es�ial. referencia al 1:288-293. Caribe. Hay dos modelos opueStos, dispersión y Guyer. C. " J. M. Savage. "1986". (1987). Cladiltic vicariancia, que intentan explicar la actual relarionlhip. amona anolea (Saoria : I.uanidae). distribución: Celestus en M6xico, América Systemati.cZobIo,y 35:509-53 1. Centr�l, Jamaica, .. Islas Caimán y Espafiola; Hedgu, S.B. 1989. Bvoluti.on and bioaeoaraphy otW'e.t Diploglosus en el:.sm'.Qe América Central, Sur Indian frogl of &he aenu. E" IÚIa.rodac" 'N8: 110\\'· América, Cuba, Puerto Rico, Monserrat y evolvina' loci and &he major ,roupI. 111: Biopoaftpli, Malpelo, y sus parientes Sauresia y Wetmorena of the Well Indiet. C.A. Wóoda (ed.)..'. Sandhlll Pre'" en la Espaftol.a. Se muestran �ladogramas GainelviUe, Florida. geogtáficos y las relaciones filogeriéticas predicbas para las formas circumcaribeftas con Leviton, A. B •• R. H. Gibb, l'R.. B. Heal • C. B. DaWIOil. base en ambos modelos. No hay aún suficiente '1985. Standard. in herpetOloáY and ichthyolo&Y: Pal't L informaclÓJi para decidirse por alguno, aunqueel Standard .ymboHc codel for inllituljOllal .... outce co1lectionl in herpetolol)' Ind ich&hyoloay. Copela vicariante es másparsim onioso. 1985:802-832. Lieberman, S. S. 1986. Bcolol), of the leaf liner a Neotrcpical rainforel'l Selva. REFERENCES herptofauna of La . . Colla Rica. 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