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TWO NEW SPECIES OF ANOLES FORMERLY REFERRED TO AS ANOLIS LIMIFRONS(SQUAMATA: POLYCHROTIDAE)

GUNTHER KO HLER1,3

AND JAVIER SUNYER1,2

1Forschungsinstitut und Naturmuseum Senckenberg, Senckenberganlage 25, 60325 Frankfurt a.M., Germany2Gabinete de Ecolog a y Medio Ambiente, Departamento de Biolog a, UNAN-Leo n, Nicaragua

ABSTRACT: We describe two new species of anoles (genusAnolis) from Panama formerly referred to as Anolis(or Norops) limifrons. Both new species differ fromA. limifronsby having a large bilobed hemipenis(small and unilobed in A. limifrons). The new species differ from each other in male dewlap size andcoloration. We provide an identification key and standardized descriptions of A. limifronsand the two new species described herein.

RESUMEN : Describimos dos especies nuevas de anoli (ge nero Anolis) de Panama anteriormente referidascomo Anolis (o Norops) limifrons. Ambas nuevas especies difieren deA. limifronsen tener un hemipene

grande y bilobulado (pequen o y unilobulado enA. limifrons). Las dos especies nuevas difieren entre s en eltamano y coloracio n de la papera gular de los machos adultos. Se proporciona una clave dicoto mica deidentificacio n y descripciones estandar de A. limifronsy de las otras dos nuevas especies.

Key words: Anolis; New species; Panama; Polychrotidae; Reptilia; Squamata

ANOLES are a diverse and taxonomically poorly understood group of lizards that aredistributed widely across the tropical andsubtropical portions of North, Central andSouth America (Ko hler, 2003; Savage, 2002).Several new species have been described inrecent years indicating that more field work and study of museum material is needed todocument the real diversity of anoles. Thecomparative study of hemipenial variation inanoles revealed substantial cryptic diversity (Ko hler and Kreutz, 1999; Ko hler et al., 2003).

In 1862, Cope described the new species Anolis (Dracontura) limifrons based on twosyntypes (now ANSP 790001) from Vera-gua. According to Savage (1970:279), today the old Veragua comprises the Provincias of Veraguas, Chiriqu and Bocas del Toro.Barbour (1934:139) described the type locality of A. limifrons as Cucuyos, Veragua Prov.,Panama [an abandoned mine on the R oSantiago]. In 1871, Cope named Anolis trochilus based on an adult male specimen(now ANSP 7804) from San Jose , CostaRica. Peters (1873) added another nominalspecies, Anolis pulchripes, based on a speci-men (now ZMB 7827) from Chiriqu . A yearlater, Cope (1874) described the new species Anolis bransfordii from Nicaragua based on

an adult male specimen (now ANSP 7890).According to Savage (1973:36), the holotypeof A. bransfordii was collected by Bransfordat Machuco (5 Machuca) on the R o SanJuan, Departamento R o San Juan, Nicara-gua. In 1882, Thominot described Anolisrivieri based on a juvenile specimen (now MNHN 1884.221) from Panama. Boulenger(1885) described Anolis godmani based onfour specimens from Guatemala and threespecimens from Irazu , Costa Rica. Stuart(1955:30) presumed that in Anolis godmaniBoulenger [there] may have been a mixupin locality data in the Godman-Salvin collec-tions, one of the cotypes having been listed asof Guatemala whereas it probably came fromCosta Rica, and he also stated that thematerial [including the holotype of A. godmani] may have been received by the BritishMuseum somewhat after the main bulk of theearlier parts of the collection had been turnedover to the Museum. In 1956, Taylordescribed the new species Anolis biscutiger based on an adult male (KU 40771) fromGolfito, Puntarenas Province, Costa Rica.Dunn (1930) regarded rivieri, trochilus and bransfordii as synonyms of limifrons. Barbour(1934:140) stated that Dunn has seen thetypes of limifrons, rivieri, pulchripes androdriguezii and declares them all the samespecies. The names trochilus, pulchripes,3 CORRESPONDENCE : e-mail, [email protected]

Herpetologica, 64(1), 2008, 92108E 2008 by The Herpetologists League, Inc.

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bransfordii and rivieri have remained in thesynonymy of A. limifronsCope whereas godmani has been retained as a valid species untilrecently (Peters and Donoso-Barros, 1970;

Savage and Villa, 1986; Taylor, 1956; Villa etal., 1988). Savage (2002) and Ko hler (2003)recognized a northern species, rodriguezii,and a southern one, limifrons, with biscutiger and godmani as synonyms of limifrons.

Here we report the results of our study of the variation in hemipenial and scalationmorphology as well as morphometrics of thesmall anoles occurring from eastern Hondurasto eastern Panama commonly referred to as Anolis (or Norops) limifrons.

MATERIALS AND METHODSFor this study, we examined 1428 specimens

of Anolis limifrons. We provide a list of thecomparative specimens examined in the Ap-pendix. Abbreviations for museum collectionsfollow those of Leviton et al., (1985) exceptMHCH (Museo Herpetolo gico de Chiriqu ,David, Panama ), and JS field numbers, whichrefer to specimens that will be deposited in theMuseo de Ciencias Naturales de la Universidad

Centroamericana (UCA), Managua, Nicara-gua. In the course of this study, G.K. had theprivilege of examining all extant primary typesof nominal species regarded as synonyms of Anolis limifronsby previous authors (Ko hler,2003; Peters and Donoso-Barros, 1970; Savage,2002). For the synonymy lists, we included only those works that cite actual specimens. No-menclature of scale characters follows that of Ko hler (2003). Terminology for hemipenialmorphology follows that of Myers et al. (1993)and Savage (1997). We everted the hemipenesin the field after euthanasia and beforepreserving the lizards in 70% ethanol by manually applying pressure to the ventral baseof tail. Once the hemipenes had been evertedto some extent, we injected 70% ethanol in thearea of the hemipenial pockets, usually resulting in complete eversion of both hemi-penes. With the needle still attached to thelizards tail, we placed the lizard for 12 mininto the preservative to allow fixation of theeverted hemipenis. We did not tie off thehemipenes at the base because this might causedamage to these fragile organs. We intended toevert the hemipenes in all available adult males

in order to document any variation. Wemeasured scale sizes using the ocular micro-meter of a stereo microscope (Leica MZ 12)and rounded the values to the nearest

0.01 mm. For all other measurements we usedprecision calipers and rounded the values tothe nearest 0.1 mm. We measured head lengthfrom tip of snout to the anterior margin of theear opening. We measured snout length fromtip of snout to the anterior border of the orbit. We determined head width as the distancebetween rictuses. We counted dorsal and ventral scales at midbody along the midline. We measured tail height and width at the pointreached by the heel of the extended hind leg. We counted subdigital lamellae on phalanges iito iv of the 4th toe. We considered the scaledirectly anterior to the circumnasal to be aprenasal. The capitalized colors and colorcodes (the latter in parentheses) are those of Smithe (19751981). To measure dewlap area, we took photographs of males with theirdewlaps artificially extended using small fore-ceps. The head portion was magnified andprinted, then superimposed on millimetricpaper, and the total number of 1-mm squarescontained in the extended dewlap counted. Astraight line was drawn between both theanterior and posterior insertions of the dewlap. We also determined the HL on the printout. We used the following equation to convert themagnified dewlap area to the real size:X 5[(! Y / A) ? B]2; where: X is the real area of thedewlap in mm2; Y is the total area (mm2) of thedewlap at a magnified scale;A is the HLmeasure (mm) of the anole at a magnifiedscale; and B is the HL measure (mm) of theanole at the real size. We executed statistical

techniques using the computer program Sta-tistica version 6.1. Juveniles were not includedin the statistical analyses. We used discrimantfunction analysis to evaluate the pheneticdistinctness of a priori groups. Discriminantscores (DC) were calculated by multiplyingselected variables by their associated canonicalcoefficients. Each specimen was then plottedalong the axae providing maximal separation of the a priori groups.

RESULTSTwo distinctly different hemipenial mor-

photypes are evident in the specimens we

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examined. In Type A ( n 5 188 adult males with everted hemipenes), the hemipenis is arelatively large bilobed organ; both thetruncus and the lobes have a strongly calycu-

late surface. The sulcus spermaticus bifurcatesat the base of the apex and the branchescontinue to the tip of the lobes. In Type B ( n5 85 adult males with everted hemipenes),the hemipenis is much smaller relative to body size as compared to the Type A hemipenis.Also, it is unilobed and the truncus and lobesare not calyculate. The sulcus spermaticusopens at the base of the apex. While these twohemipenial types show a broadly sympatricgeographical distribution pattern in westernPanama, variation in hemipenial morphology within these discrete types is negligible both within populations and in a geographicalcontext. Within Type A, two distinct typescan be distinguished in respect of the relativesize and the coloration of the male dewlap.Males with bilobed hemipenes from theProvince of Bocas del Toro, Panama, andadjacent southeastern Costa Rica have a smalldewlap (smaller than 100 mm2) that is dull white with a small basal orange blotch. Males with bilobed hemipenes from central andeastern Panama have a large dewlap (largerthan 150 mm2) that is almost uniformly orange. All Type B males have a small dewlap(smaller than 100 mm2) that is dull white witha small basal orange blotch.

In external morphology, there is greatoverlap in the ranges of all examined charac-ters of scalation and morphometrics (Table 1). We conducted a discriminant function analy-sis (DFA) based on seven pholidotic charac-ters (number of medial ventral scales in onehead length; number of medial dorsal scales inone head length; subdigital lamellae; totalnumber of loreal scales; scales betweensupraorbital semicircles; scales between pos-terior canthals; scales around midbody) anddid a priori assignments to groups based onour hemipenis and male dewlap findings(Group 1 5 Type B hemipenis, male dewlapsmall, dull white with a small basal orangeblotch; Group 2 5 Type A hemipenis, maledewlap small, dull white with a small basalorange blotch; Group 3 5 Type A hemipenis,male dewlap large, more or less uniformly orange). This DFA yielded a scatter diagram

(Fig. 1) that correctly classified 72.7% of thespecimens. The first function is DS5 0.471297(number of medial ventral scales in one headlength) + 0.673627 (number of medial dorsal

scales in one head length)2

0.392336 (sub-digital lamellae) + 0.176923 (total number of loreal scales) + 0.286056 (scales between su-praorbital semicircles) 2 0.651674 (scales be-tween posterior canthals) + 0.025958 (scalesaround midbody). The second function is DS5 0.284551 (number of medial ventral scales inone head length) 2 0.173192 (number of medial dorsal scales in one head length)2 0.610593 (subdigital lamellae) 2 0.171498(total number of loreal scales) 2 0.293526(scales between supraorbital semicircles)+ 0.054329 (scales between posterior canthals)2 0.603442 (scales around midbody).

Based on our data, we recognize threespecies of this complex: Species A: Hemipenisunilobed; male dewlap small (smaller than100 mm2), dull white with a small basalorange blotch; distributed from eastern Hon-duras to central Panama west of the CanalZone (Fig. 2). Species B: Hemipenis bilobed;male dewlap small (smaller than 100 mm2),dull white with a small basal orange blotch;distributed in the western and central portionsof the Province Bocas del Toro, Panama, andadjacent southeastern Costa Rica (Fig. 2).Species C: Hemipenis bilobed; male dewlaplarge (larger than 150 mm2), almost uniformly orange; distributed in central and easternPanama; expected in adjacent northwesternColombia (Fig. 2). In western Panama (Prov-inces of Chiriqu , Bocas del Toro and Veraguas), the bilobed form (our Species B)is restricted to the Caribbean versant; whereasthe unilobed form (our Species A) occurs onboth versants. Although no cases of actualsyntopy of these two forms have beendocumented, they show a broadly sympatricdistribution pattern on the Caribbean versantof western Panama.

Unfortunately, none of the type specimensmentioned in the introduction are males withtheir hemipenes everted. Also, data on dewlapcoloration in life is not available for these typespecimens. However, the type locality data of most of these nominal species allow for areliable allocation to one of the three speciesthat we distinguish. Thus, on geographic

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reasons the following taxa (respective typelocalities in parentheses) are clearly referableto our Species A: Anolis limifrons Cope(Cucuyos, Veragua Province, Panama); A. trochilus Cope (San Jose , Costa Rica); A. pulchripes Peters (Chiriqu , Panama); A. bransfordii Cope (Nicaragua); A. godmani:Boulenger (Irazu , Costa Rica); andA. biscutiger Taylor (Golfito, Puntarenas Province, CostaRica). Because A. limifronsCope is the oldestavailable name for this species, our Species Ahas to be referred to that name and the othernominal species remain in the synonymy of A.limifrons. The holotype of A. rivieri Thominotis a juvenile with unspecific locality data(Panama), and its taxonomic identity cannotbe determined. Therefore, we consider A.rivieri Thominot to be a nomen dubium.Interestingly, there is no available scientificname for either of our Species B and C. We

therefore describe them as new species below. Anolis limifronsCope, 1862

Anolis biscutiger Taylor, 1956:81. HolotypeKU 40771 from Golfito, Puntarenas Prov-ince, Costa Rica.

Anolis bransfordii Cope, 1874:67. HolotypeANSP 7890 from Nicaragua.

Anolis godmaniBoulenger, 1885:85. Syntypes(BMNH and MCZ) from Guatemala andIrazu, Costa Rica. Barbour (1834), Taylor(1956).

Anolis limifronsCope, 1862:178. SyntypesANSP 790001 from Veragua. Bocourt

(1873; in part); Dunn (1930; in part);Barbour (1934; in part); Gaige et al.,(1937); Brattstrom and Howell (1954);Taylor (1956; in part); Meyer and Wilson

(1973); Fitch and Seigel (1984); Vences etal., (1998); Poe (2004; in part). Anolis limifrons limifrons: Etheridge (1959; in

part). Anolis pulchripesPeters, 1873:739. Holotype

ZMB 7827 from Chiriqui. Anolis trochilus Cope, 1871:215. Holotype

ANSP 7804 from San Jose , Costa Rica.Cope (1876); Savage (1970).

Norops limifrons: Wilson and McCranie(1994); Ko hler (1999); Ko hler (2001); Ko h-

ler et al., (2001); Ko

hler and McCranie(2001).Diagnosis.A medium-sized species

(snoutvent length [SVL] in largest specimen43.5 mm) of the genus Anolis (sensu Poe,2004) that is most similar in external mor-phology to a cluster of Central Americanspecies that are long-legged (longest toe of adpressed hindlimb reaches to mid-eye orbeyond), have a single elongated prenasalscale, smooth to slightly keeled ventral scales,

and slender habitus, often delicate (i.e.,Anolisdollfusianus, A. ocelloscapularis, A. rodriguezii, A. yoroensis, A. zeus). Within this clusterof species, A. limifrons can be readily distinguished by male dewlap coloration (dull white with a small basal orange blotch inA.limifrons vs. uniformly dull white inA. zeus,and almost uniformly orange to orange-yellow in the remaining species. Additionally, A.limifrons differs from the species in thiscluster by the following characteristics (con-dition for A. limifronsin parentheses): Anolisdollfusianus: ventrals weakly keeled (smooth). Anolis ocelloscapularis: An ocellated shoulderspot present (absent); ventrals weakly keeled(smooth); hemipenis bilobed (unilobed).Ano-lis rodriguezii: hemipenis bilobed (unilobed). Anolis yoroensis: ventrals weakly keeled(smooth).

Description (Fig. 3).Maximum SVL 41.5mm in males, 43.5 mm in females; ratio taillength/SVL 1.532.52 (2.17 6 0.21); tailslightly compressed in cross section, ratio tailheight/tail width 1.001.30 (1.166 0.07); ratioaxilla to groin distance/SVL 0.360.53 (0.4360.03); ratio head length/SVL 0.240.28 (0.26

FIG. 1.Discriminant function analysis of the CentralAmerican anoles formerly referred to asAnolis limifrons.See text for details.


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6 0.01); ratio snout length/head length 0.410.52 (0.45 6 0.02); ratio head length/head width 1.441.80 (1.676 0.06); longest toe of adpressed hind limb reaching to a pointbetween anterior to eye and tip of snout; ratioshank length/SVL 0.260.33 (0.296 0.02);ratio shank length/head length 0.971.24 (1.116 0.07); longest finger of extended forelimbreaching to a point between nostrils and tip of snout; longest finger of adpressed forelimbreaches in between anterior to insertion of hind limbs and slightly beyond to insertion of

hind limbs; scales on snout varying fromalmost non-keeled to keeled; 59 (6.96 0.8)postrostrals; 711 (8.9 6 0.9) scales betweennasals; 1 large elongated prenasal scale incontact with both rostral and first supralabial,occasionally only in contact with rostral; scalesin distinct prefrontal depression generally slightly tuberculate posteriorly, wrinkled an-teriorly, some of them keeled; supraorbitalsemicircles well developed, separated by 04(1.8 6 0.8) scales; supraorbital disc composedof 514 distinctly enlarged keeled scales;

FIG. 2.Map indicating known collecting sites mentioned in text of Anolis apletophallus (triangles), A.cryptolimifrons (squares), and A. limifrons (circles) in Honduras, Nicaragua, Costa Rica and Panama. Each symbol

can represent one or more nearby localities. Areas above 500 and 1000 m are shaded gray. Open symbols: specimens with no everted hemipenis; Solid symbols: specimens with everted hemipenis.


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circumorbital row usually incomplete, there-fore, 03 enlarged supraorbitals in contact with supraorbital semicircles; a single largeelongated superciliary; 36 rows of smallkeeled scales extending between enlargedsupraorbitals and superciliaries; a very shallow parietal depression present in most speci-mens; interparietal scale well developed,usually surrounded by scales of moderate sizeanteriorly and by small to moderate size scalesposteriorly; 15 (2.7 6 0.9) scales presentbetween interparietal and supraorbital semi-circles; canthal ridge distinct, composed of 611 (7.5 6 0.8) canthal scales, with 35 (4.060.7) larger posterior scales; 715 (10.66 1.7)scales present between second canthals; 1018 (13.8 6 2.1) scales present betweenposterior canthals; 2468 (41.16 9.6) lorealscales in a maximum of 58 (6.16 0.8)horizontal rows, with the scales of lower rowsand those adjacent to the canthals mostly keeled, and those of upper rows non-adjacent

to the canthals mostly tuberculated; 47keeled subocular scales arranged in a singlerow; 58 (6.5 6 0.7) supralabials to levelbelow center of eye; 25 suboculars broadly in

contact with supralabials; ear opening medi-um-sized, ratio tympanum height/interparietalscale length 0.581.36 (0.866 0.16); mentaldistinctly wider than long, completely dividedmedially, bordered posteriorly by 48 (6.560.9) postmentals; 58 (6.36 0.7) infralabialsto level below center of eye; sublabialsundifferentiated; keeled granular scales pre-sent on chin and throat; dewlap extendingfrom level below oral ricti to axilla, in somespecimens extending 12 mm posterior to

axilla; dorsum of body with weakly keeledgranular scales (at least anteriorly) withrounded posterior margins, 2 medial rowsslightly enlarged, 3870 (54.36 5.8) medialdorsal scales in one head length; 70112 (91.56 9.0) medial dorsal scales between axilla andgroin; lateral scales homogeneous, ventrals atmidbody smooth, slightly bulging, nonimbri-cate, 2658 (40.6 6 6.1) ventral scales in onehead length; 4979 (64.66 5.7) ventral scalesbetween axilla and groin; 108157 (132.1610.6) scales around midbody; caudal scalesstrongly keeled; caudal middorsal scalesslightly enlarged, without whorls of enlargedscales, although an indistinct division insegments is discernible; a pair of slightly enlarged postanal scales usually present; notube-like axillary pocket present; scales ondorsal surface of forelimb keeled, imbricate;digital pads dilated; distal phalanx narrowerthan and raised from dilated pad; 2027 (23.36 1.6) lamellae under phalanges iiiv of fourth toe; 711 (8.5 6 0.9) scales underdistal phalanx of fourth toe.

Description of completely everted hemipenis.Small unilobed organ; sulcus spermati-cus bordered by well developed sulcal lips andopens at base of apex; no discernable surfacestructure on truncus and lobes; no asulcateprocessus present (Fig. 4).

Anolis apletophallus sp. nov. Anolis limifrons: Bocourt (1873; in part);

Dunn (1930; in part); Barbour (1934; inpart); Breder (1946).

Anolis limifrons limifrons: Etheridge (1959; inpart).

FIG. 3.Head of Anolis limifrons(SMF 86900). Scalebars equal 1.0 mm.


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Holotype (Fig. 5).SMF 85307, an adultmale from Panama City, Metropolitan Na-tional Park (8u 589600N, 79u 329460 W), 45 m,Panama Province, Panama. Collected 26January 2006 by Gunther Ko hler, JavierSunyer, Abel A. Batista R. and Marcos Ponce.Field tag number GK 1672.

Paratypes.SMF 8530819, same collect-ing data as holotype. SMF 8530813 are adultmales, SMF 8531419 are adult females.

Diagnosis.A medium-sized species (SVL

in largest specimen 47.0 mm) of the genus Anolis(sensu Poe, 2004) that is most similar inexternal morphology to a cluster of CentralAmerican species that are long-legged (lon-gest toe of adpressed hindlimb reaches tomid-eye or beyond), have a single elongatedprenasal scale, smooth to slightly keeled ventral scales, and slender habitus, oftendelicate (i.e., Anolis dollfusianus, A. limifrons, A. ocelloscapularis, A. rodriguezii, A. yoroensis, A. zeus). Within this cluster of species,A.apletophallus is most similar to A. limifronsfrom which it is readily distinguished by hemipenis morphology: hemipenis small andunilobed in A. limifrons, large and bilobed in A. apletophallus(Fig. 6). Anolis apletophallusdiffers from the remaining species in thiscluster by the following characteristics (con-dition for A. apletophallus in parentheses): Anolis dollfusianus: Hemipenis unilobed (bi-lobed); ventrals weakly keeled and slightly imbricate (smooth and non-imbricate).Anolisocelloscapularis: Ventrals weakly keeled andslightly imbricate (smooth and non-imbricate);an ocellated shoulder spot present (absent). Anolis rodriguezii: Ventrals weakly keeled and

slightly imbricate (smooth and non-imbricate). Anolis yoroensis: Ventrals weakly keeled andslightly imbricate (smooth and non-imbricate). Anolis zeus: Hemipenis unilobed (bilobed);male dewlap uniformly dull white (almostuniformly orange).

Description of the holotype.Adult male asindicated by everted hemipenes; SVL44.0 mm; tail length 96.0 mm, tail complete;tail slightly compressed in cross section, tailheight 1.75 mm, tail width 1.70 mm; axilla togroin distance 17.6 mm; head length 12.0 mm,head length/SVL ratio 0.27; snout length5.2 mm; head width 7.1 mm; longest toe of adpressed hind limb reaching to a pointbetween eyes and rostrils; shank length13.0 mm, shank length/head length ratio 1.08;longest finger of extended forelimb reaching toa point slightly beyond nostrils; longest fingerof adpressed forelimb just reaches anteriorinsertion of hind limbs. Most scales on snout

FIG. 5.Head of holotype of Anolis apletophallus(SMF 85307). Scale bars equal 1.0 mm.

FIG. 4.Hemipenisof Anolis limifrons(SMF 85246). (a)sulcate view; (b) asulcate view. Scale bar equals 1.0 mm.


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keeled; 7 postrostrals; 10 scales between nasals;1 large elongated prenasal scale in contact withboth rostral and first supralabial; scales indistinct prefrontal depression slightly tubercu-late posteriorly, wrinkled anteriorly; supraor-bital semicircles well developed, separated by 3scales; supraorbital disc composed of 1315distinctly enlarged keeled scales; circumorbitalrow incomplete, therefore, one enlarged su-praorbital in contact with supraorbital semicir-cles; a single large elongated superciliary; about3 or 4 rows of small keeled scales extendingbetween enlarged supraorbitals and super-ciliaries; a very shallow parietal depressionpresent; interparietal scale well developed, 1.43 1.0 mm (length 3 width), surrounded by scales of moderate size; 3 scales presentbetween interparietal and supraorbital semi-circles; canthal ridge distinct, composed of 4large (posterior two largest) and 4 smallanterior canthal scales; 10 scales presentbetween second canthals; 17 scales present

between posterior canthals; 64 (right)61 (left)loreal scales in a maximum of 8 horizontal rows, with the scales of lower rows mostly keeled,and those of upper rows mostly tuberculated; 8

(right)6 (left) keeled subocular scales ar-ranged in a single row; 7 supralabials to levelbelow center of eye; 4 (right)3 (left) subocu-lars broadly in contact with supralabials; earopening 0.90 3 1.40 mm (length 3 height);mental distinctly wider than long, completely divided medially, bordered posteriorly by keeled 6 postmentals (outer pair larger); 7infralabials to level below center of eye;sublabials undifferentiated; keeled granularscales present on chin and throat; dewlapextending from level below oral ricti to 3 mmbeyond level of axilla; dorsum of body with weakly keeled granular scales with roundedposterior margins, 2 medial rows slightly enlarged, largest dorsal scales about 0.2130.23 mm (length 3 width); about 64 medialdorsal scales in one head length; about 85medial dorsal scales between axilla and groin;lateral scales homogeneous, average size0.16 mm in diameter; ventrals at midbody smooth, slightly bulging, non-imbricate, about0.34 3 0.34 mm (length 3 width); about 42 ventral scales in one head length; about 66 ventral scales between axilla and groin; 142scales around midbody; caudal scales strongly keeled; caudal middorsal scales slightly en-larged, without whorls of enlarged scales,although an indistinct division in segments isdiscernible; a pair of enlarged postanal scalespresent, about 0.69 mm wide; no tube-likeaxillary pocket present; scales on dorsal surfaceof forelimb keeled, imbricate, about 0.28 30.25 mm (length3 width); digital pads dilated;distal phalanx narrower than and raised fromdilated pad; 24 lamellae under phalanges iiiv of fourth toe; 9 (right)8 (left) scales underdistal phalanx of fourth toe.

Description of completely everted hemipenis.Medium-sized bilobed organ; sulcusspermaticus bordered by well developed sulcallips and bifurcating at base of apex; shortly afterthe bifurcation, branches open into broadconcave areas, one on each lobe; asulcatesurface of apex and distal truncus strongly calyculate, base of truncus with transversefolds; no asulcate processus present, althougha slightly elevated ridge present (Fig. 6).

FIG. 6.Hemipenis of Anolis apletophallus (SMF80719). (a) sulcate view; (b) asulcate view. Scale barequals 1.0 mm.


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Coloration in life.Dorsal ground color Warm sepia (color 221A); flanks Dark Drab(119B); dorsal surface of head Dark Drab(119B); venter dull white suffused with Dark

Drab (119B); dorsal surface of limbs MarsBrown (223A); tail Drab Gray (119D) withtransverse Sepia (119) (but slightly morereddish) bands; iris Raw Sienna (136); dewlapOrange Yellow (18).

Variation.The paratypes agrees well withthe holotype in general appearance, morpho-metrics and scalation (see Table 1). In most of the male paratypes, one pair of slightly enlarged postanal scales is present. Thefemale paratypes have no dewlap and noenlarged postanal scales. The coloration in lifeof male paratypes dewlap (SMF 85308) wasrecorded as Trogon Yellow (153).

Etymology.The name apletophallus isformed from the Greek words apletos (im-mense) and phallos (penis) and is used as anoun in apposition.

Natural history notes.All type specimens were collected active during the day insecondary forest. The lizards were spottedon tree trunks and fallen branches, from nearground level to 1.5 m above the ground.

Anoliscryptolimifrons sp. nov.

Anolis limifrons: Poe (2004; in part). Anolis limifrons limifrons: Etheridge (1959; in

part.).

Holotype (Fig. 7).SMF 85230, an adultmale from Cerro Brujo (9u 11916.40N,82u 11925.40 W), 10 m, Bocas del Toro Prov-ince, Panama. Collected 19 January 2006 by Gunther Ko hler, Javier Sunyer, Abel A.Batista R. and Marcos Ponce. Field tagnumber GK 1502.

Paratypes.SMF 8523144, same collect-ing data as holotype. Most of the paratypes arefemales, except SMF 8523637, 8523940,8524243 (adult males).

Diagnosis.A medium-sized species (SVLin largest specimen 45.0 mm) of the genus Anolis(sensu Poe, 2004) that is most similar inexternal morphology to a cluster of CentralAmerican species that are long-legged (lon-gest toe of adpressed hindlimb reaches tomid-eye or beyond), have a single elongatedprenasal scale, smooth to slightly keeled

ventral scales, and slender habitus, oftendelicate (i.e., A. apletophallus, A. dollfusianus, A. limifrons, A. ocelloscapularis, A. rodriguezii, A. yoroensis, A. zeus). Within this clusterof species,A. cryptolimifronsis most similar to A. limifrons and A. apletophallus. Anoliscryptolimifrons is readily distinguished from A. limifronsby hemipenis morphology: hemi-penis small and unilobed inA. limifrons, largeand bilobed in A. cryptolimifrons. Anoliscryptolimifrons differs from A. apletophallusin male dewlap size and color in life (small anddull white with a small basal orange blotch inN. cryptolimifrons versus dewlap large andalmost uniformly orange inN. apletophallus). Anolis cryptolimifronsdiffers from the re-maining species in this cluster by the followingcharacteristics (condition for A. cryptolimifrons in parentheses): Anolis dollfusianus:Hemipenis unilobed (bilobed); ventrals weak-ly keeled (smooth); fewer than 80 dorsals

FIG. 7.Head of holotype of Anolis cryptolimifrons(SMF 85230). Scale bars equal 1.0 mm.


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between levels of axilla and groin (more than87); male dewlap almost uniformly orange- yellow (dull white with a small basal orangeblotch). Anolis ocelloscapularis: An ocellated

shoulder spot present (absent); ventrals weak-ly keeled (smooth); male dewlap almostuniformly orange (dull white with a smallbasal orange blotch).Anolis rodriguezii: Maledewlap almost uniformly orange (dull white with a small basal orange blotch).Anolis yoroensis: Ventrals weakly keeled (smooth);fewer than 87 dorsals between levels of axillaand groin (more than 87); male dewlap almostuniformly orange (dull white with a smallbasal orange blotch). Anolis zeus: Hemipenisunilobed (bilobed); male dewlap uniformly dull white without a basal orange blotch (basalorange blotch present).

Description of the holotype.Adult male asindicated by everted hemipenes; SVL41.0 mm; tail length 88.0 mm, tail complete;tail slightly compressed in cross section, tailheight 1.50 mm, tail width 1.25 mm; axilla togroin distance 18.0 mm; head length11.0 mm, head length/SVL ratio 0.27; snoutlength 5.2 mm; head width 6.6 mm; longesttoe of adpressed hind limb reaching toanterior portion of eye; shank length11.1 mm, shank length/head length ratio1.01; longest finger of extended forelimbreaching nostrils; longest finger of adpressedforelimb not reaching anterior insertion of hind limbs. Most scales on snout keeled, some wrinkled; 8 postrostrals; 10 scales betweennasals; 1 large elongated prenasal scale incontact with both rostral and first supralabial;scales in distinct prefrontal depression slightly tuberculate; supraorbital semicircles well developed, separated by 2 scales; supraorbitaldisc composed of 910 distinctly enlargedkeeled scales; circumorbital row complete,therefore, no enlarged supraorbitals in contact with supraorbital semicircles; a single largeelongated superciliary; about 5 rows of smallkeeled scales extending between enlargedsupraorbitals and superciliaries; a shallow parietal depression present; interparietal scale well developed, 1.303 0.95 mm (length 3 width), surrounded by scales of moderate sizeanteriorly and small size posteriorly; 3 scalespresent between interparietal and supraorbitalsemicircles; canthal ridge distinct, composed

of 4 large (posterior two largest) and 4 smallanterior canthal scales; 16 scales presentbetween second canthals; 19 scales presentbetween posterior canthals; 61 loreal scales in

a maximum of 6 horizontal rows, with thescales of lower rows mostly keeled, and thoseof upper rows mostly tuberculated; 6 (right) 5 (left) keeled subocular scales arranged in asingle row; 7 supralabials to level below centerof eye; 2 suboculars broadly in contact withsupralabials; ear opening 0.5 3 1.3 mm(length 3 height); mental distinctly widerthan long, completely divided medially, bor-dered posteriorly by 8 keeled postmentals(outer pair larger); 7 infralabials to level below center of eye; sublabials undifferentiated;keeled granular scales present on chin andthroat; dewlap extending from level below oralricti to about 1 mm anterior to axilla; dorsumof body with weakly keeled granular scales with rounded posterior margins, 2 medialrows slightly enlarged, largest dorsal scalesabout 0.19 3 0.20 mm (length 3 width);about 54 medial dorsal scales in one headlength; about 97 medial dorsal scales betweenaxilla and groin; lateral scales homogeneous,average size 0.10 mm in diameter; ventrals atmidbody smooth, slightly bulging, non-imbri-cate, about 0.24 3 0.27 mm (length 3 width);about 46 ventral scales in one head length;about 68 ventral scales between axilla andgroin; 148 scales around midbody; caudalscales strongly keeled; caudal middorsal scalesslightly enlarged, without whorls of enlargedscales, although an indistinct division insegments is discernible; a pair of enlargedpostanal scales present; no tube-like axillary pocket present; scales on dorsal surface of forelimb keeled, imbricate, about 0.24 30.26 mm (length 3 width); digital padsdilated; distal phalanx narrower than andraised from dilated pad; 24 lamellae underphalanges iiiv of fourth toe on right foot(fourth toe of left foot missing); 7 scales underdistal phalanx of fourth toe.

Description of completely everted hemipenis.Medium-sized bilobed organ; sulcusspermaticus bordered by well developed sulcallips and bifurcating at base of apex; shortly afterthe bifurcation, branches open into broadconcave areas, one on each lobe; asulcatesurface of apex and distal truncus strongly


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calyculate, base of truncus with transversefolds; no asulcate processus, although a slightly elevated ridge present (Fig. 8).

Coloration in life.Dorsal ground colorRaw Umber (color 123 in Smithe 19751981)suffused with Vandyke Brown (221) at mid-dorsum; venter Pale Horn Color (92); irisCinnamon (123A); dewlap dull white with asmall Chamois (123D) basal blotch.

Variation.The paratypes agrees well withthe holotype in general appearance, morpho-metrics and scalation (see Table 1). In most of the male paratypes, one pair of slightly enlarged postanal scales is present. Thefemale paratypes have no dewlap and noenlarged postanal scales.

Etymology.The species name cryptolimifrons is used as a noun in apposition andreflects the similarity and suspected close

relationship between the new species and itscongener Anolis limifronsCope.

Natural history notes.All type specimens were collected active during the day in a patch

of secondary forest. The lizards were spottedon branches and leaves of bushes and smalltrees, 0.5 to 1.5 m above the ground.

Key to the males of the species formerly referred to as Anolis limifrons

1a. Male dewlap large, larger than 150 mm2

(Fig. 9a), almost uniformly orange_ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Anolis apletophallus

1b. Male dewlap small, smaller than 100 mm2

(Figs. 9b,c), dull white with a small basal

orange-yellow blotch_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _

22a. Hemipenis unilobed_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _Anolis limifrons2b. Hemipenis bilobed_ _ _ _ _ _ _ _ _Anolis cryptolimifrons

DISCUSSIONUntil recently, hemipenial morphology in

the diverse group of anoles has been mostly ignored. The minority of male anoles housedin museum collections worldwide have evert-ed hemipenes. We urge future collectors of these anoles to attempt to fully evert thehemipenes of their specimens. Study of theseorgans may reveal more cryptic species of anoles than originally envisaged by most workers. This distinction is of great conserva-tional importance: whereas a large amount of ecological data is available forAnolis limifronsand A. apletophallus, very little is known from A. cryptolimifrons, a species with a relatively restricted distribution. Although A. limifronsand A. cryptolimifronsboth occur in the samegeneral area of mainland western Panama(Caribbean versant), no actual instance of syntopy has been documented. Both taxaappear to use the same habitat and sameperching sites and might exclude each other.

Savage (2002) reported populations of Anolis limifrons-like lizards from the slopesof the volcanoes Irazu and Turrialba withkeeled ventral scales but otherwise identical inscutellation and coloration. Boulenger (1885)based the description of A. godmani on thischaracteristic, and in earlier publicationsSavage (e.g., Savage, 1973; Savage and Villa,1986) recognized this form as a valid speciesand used the name A. godmani for it. More

FIG. 8.Hemipenis of holotype of Anolis cryptolimifrons (SMF 85230). (a) sulcate view; (b) asulcate view.Scale bar equals 1.0 mm.


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recently, Savage (2002) considered it to be anindividual variant and therefore a synonym of A. limifrons. We have examined a singlefemale (SMF 86924) from this general areacollected at 1500 m elevation (about 160 mabove the highest Costarican record for this

species) which has distinctly keeled ventralscales. Also, the scales on head and dorsumare strongly keeled, much more than in any other examined A. limifrons. Unfortunately,

no data on hemipenis morpholgy nor on maledewlap coloration are available for represen-tatives of this population. Further collectingand study is needed in order to evaluate thestatus of the populations of A. limifrons-likelizards from the slopes of the vulcanos Irazu and Turrialba in Costa Rica.

Acknowledgments.Collecting and exportation permits were provided by J. Guevara Sequeira, SINAC Central,Ministerio del Ambiente y Energ a (MINAE), San Jose ,Costa Rica; A. Barahona, T. Garc a, A. P. Martinez, E.Mun oz Galeano, and C. Romero, Corporacio n Hondur-

en a de Desarollo Forestal (COHDEFOR), Tegucigalpa,Honduras; M. Fonseca Cuevas, S. Tijerino, B. Quintero, I.Ortega, M. G. Camacho, C. Mej a, and C. Peres-Roma n,Ministerio del Ambiente y los Recursos Naturales(MARENA), Managua, Nicaragua; Y. Hidalgo, AutoridadNacional del Ambiente (ANAM), Panama City, Panama.A. Batista and M. Ponce (Panama), and A. Hertz, S.Lotzkat, D. Manzanarez and L. Obando (Nicaragua),actively helped in the field. M. Dehling, J. Ferrari, G. Paizand J. R. McCranie generously donated specimens for thisstudy. M. Piepenbring, Botanisches Institut J. W. Goethe-Universita t, Frankfurt, Germany, B. E. Sanjur, Facultadde Ciencias Naturales y Exactas de la UniversidadAuto noma de Chiriqu (UNACHI), David, Panama, andJ. Vegas, Panama City, provided logistical support for ourstudies in Panama. This paper is based in part upon work supported by the Deutscher Akademischer Austausch-dienst (DAAD) to J. Sunyer and to G. Ko hler through thePartnership Program between the J. W. Goethe-Universi-ta t Frankfurt am Main, Germany, und der Facultad deCiencias Naturales y Exactas der Universidad Auto nomade Chiriqu (UNACHI), David, Panama. We thank L.Czupalla and J. Ko hler for providing some of the drawingsused in this paper. For the loan of and/or access tospecimens, we thank L. Ford, C. J. Raxworthy and D. R.Frost, American Museum of Natural History (AMNH),New York; T. Daeschler and A. Gilmore, Academy of Natural Sciences (ANSP), Philadelphia; S. P. Rogers,Carnegie Museum of Natural History (CM), Pittsburgh;A. Resetar, Field Museum of Natural History (FMNH),Chicago; G. Lenglet, Institut Royal des Sciences Natur-elles de Belgique (IRSNB), Bruxelles; W. E. Duellmanand J. E. Simmons, University of Kansas, Natural History Museum (KU), Lawrence; D. Rossman, Museum of Natural Science, Louisianna State University (LSUMZ),Baton Rouge; J. Hanken and J. P. Rosado, Museum of Comparative Zoology, Harvard University (MCZ), Cam-bridge; A. Batista and M. Ponce, Museo Herpetolo gico deChiriqu (MHCH), David; F. Tiedemann, Naturhistor-isches Museum (NMW), Wien; F. Bolan os, Museo deZoolog a Universidad de Costa Rica (UCR), San Jose ; K.

L. Krysko and F. W. King, Florida Museum of NaturalHistory (UF), Gainesville; C. A. Phillips and J. Petzing,Illinois Natural History Survey, Center for Biodiversity (UIMNH), Champaign; R. A. Nussbaum and G. Schnei-

FIG. 9.Specimens in life with extended dewlaps (a) Anolis apletophallusmale (Panama City, Panama , Panama;not preserved); (b) A. cryptolimifronsmale (Cerro Brujo,Bocas del Toro, Panama; not preserved); (c)A. limifrons,male (Los Algarrobos, Chiriqu , Panama; not preserved).


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der, University of Michigan Museum of Zoology (UMMZ), Ann Arbor; R. W. McDiarmid and W. R.Heyer, National Museum of Natural History (USNM), Washington, D.C.; J. Campbell and C. Franklin, TheUniversity of Texas at Arlington (UTA), Arlington; W.Bo hme, Zoologisches Forschungsinstitut und Museum A.Koenig (ZFMK), Bonn; and R. Gunther, Museum fu rNaturkunde der HumboldtUniversita t zu Berlin (ZMB),Berlin. We thank M. Harvey and two anonymous reviewersfor comments on an earlier version of the manuscript.

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COPE, E. D. 1876. On the Batrachia and Reptilia of CostaRica. Journal of the Academy of Natural Sciences of Philadelphia 8:93154. (1875)

DUNN, E. R. 1930. Notes on Central AmericanAnolis.Proceedings of the New England Zoological Club12:1524.

ETHERIDGE , R. 1959. The relationships of the anoles(Reptilia: Sauria: Iguanidae). An interpretation basedon skeletal morphology. Ph.D. Dissertation, University of Michigan, Ann Arbor, Michigan, U.S.A.

FITCH, H. S., AND R. A. SEIGEL . 1984. Ecological andtaxonomic notes on Nicaraguan anoles. Milwaukee PublicMuseum Contributions in Biology and Geology 57:113.

GAIGE, H. T., N. HARTWEG, AND L. C. STUART. 1937. Noteson a collection of amphibians and reptiles from easternNicaragua. Occasional Papers Museum of Zoology

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KO HLER , G. 2001. Anfibios y Reptiles de Nicaragua.Herpeton, Offenbach, Germany.

KO HLER , G. 2003. Reptiles of Central America. Herpeton Verlag, Offenbach, Germany.

KO HLER , G., AND J. KREUTZ. 1999. Norops macrophallus

(Werner, 1917), a valid species of anole from Guate-mala and El Salvador (Squamata: Sauria: Iguanidae).Herpetozoa 12:5765.

KO HLER , G., AND J. R. MCCRANIE. 2001. Two new speciesof anoles from northern Honduras (Squamata: Poly-chrotidae). Senckenbergiana Biologica 81:235245.

KO HLER , G., J. R. MCCRANIE, K. E. NICHOLSON , AND J.KREUTZ. 2003. Geographic variation in hemipenialmorpholgy in Norops humilis (PETERS 1863), and thesystematic status of Norops quaggulus (COPE 1885)(Reptilia, Squamata: Polychrotidae). SenckenbergianaBiologica 82:213222.

KO HLER , G., J. R. MCCRANIE, AND L. D. W ILSON. 2001. Anew species of anole from western Honduras (Squa-mata: Polychrotidae). Herpetologica 57:247255.

LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E.DAWSON. 1985. Standards in herpetology and ichthyol-ogy: Part I. Standard symbolic codes for institutionalresource collections in herpetology and ichthyology.Copeia 1985:802832.

MEYER, J. R., AND L. D. W ILSON. 1973. A distributionalchecklist of the turtles, crocodilians, and lizards of Honduras. Contributions in Science, Natural History Museum, Los Angeles County 244:139.

MYERS, C. W., E. E. W ILLIAMS, AND R. W. MCDIARMID.1993. A new anoline lizard (Phenacosaurus) from thehighland of Cerro de la Neblina, southern Venezuela.

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PETERS, W. 1873. U ber neue Saurier (Spaeriodactylus, Anolis, Phrynosoma, Tropidolepisma, Lygosoma,Ophioscincus) aus Centralamerica, Mexico und Aus-tralien. Monatsbericht der Ko niglich Preussis-chen Akademie der Wissenschaften zu Berlin 1873:738747.

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SMITHE , F. B. 19751981. Naturalists color guide. Part I.Color guide. 182 color swatches. American Museum of Natural History, New York, New York, U.S.A.

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. Accepted:9 January 2008

. Associate Editor:Michael Harvey

APPENDIX Specimens Examined.* Specimens used in Discrimi-

nant Function AnalysisH Specimens with everted hemipenis Anolis apletophallus.Panama: Colo n: Cano Saddle,

Closes plantation: USNM 69586; Chagres River: USNM102849; R o Indio, near Gatu n: USNM 54007; Gatu n:USNM 5401112; Bohio Peninsula East, Panama Canal:USNM 50523137, 50523842all H , 50524351, 505252H ,505253, 50525456all H , 505257, 505258H , 505259,505260H , 50526169, 50527073a ll H , 505274, 50527576both H , 50527785, 505286H , 50528788, 50528990both H ,50529192, 505293301all H ; Buena Vista Peninsula,Panama Canal: USNM 505199230; Gigante Ridge, Pana-ma Canal: USNM 50537182, 50538384both H , 505385,50538688all H , 50538993, 50539495both H , 50539697,50539899both H , 505400, 505401H ; Juan Gallegos Island East, Panama Canal: USNM 50557182, 50558384both H ,505585, 505586H , 50558789, 505590H , 50559197,50559899both H , 505600, 505601H ; Juan Gallegos Island North, Panama Canal: USNM 50560204, 50560508all H ,50560916, 505617H , 50561822, 505623H , 505624,505625H , 505626, 505627H ; Juan Gallegos Island South,Panama Canal: USNM 50554149, 505550H , 50555152,505553H , 505554, 50555557all H , 505558, 505559H ,50556063, 50556468all H , 505569, 505570H ; LimboCamp, Panama Canal: USNM 50545773, 505474H ,50547579, 50548081both H , 505482, 505483H ; Lion HillIsland, Panama Canal: USNM 50530205, 50530608all H ,505309, 505310H , 50531118, 50531920both H , 50532131,505332H , 50533334; Pen a Blanca Peninsula, PanamaCanal: USNM 50540206, 505407H , 50540810, 50541115all H , 505416, 505417H , 50541822, 505423H , 50542425,50542627both H , 505428, 50542930both H , 505431; Poach-

ers Peninsula: USNM 50565059, 50566061both H

,50566264, 505665H , 50566667, 50566869both H ,505670, 50567173all H , 50567476; Puma Island, PanamaCanal: USNM 50533544, 505345H , 50534650, 505351H ,

50535253, 50535456all H , 50535760, 505361H , 50536263, 505364H , 505365, 505366H , 505367, 50536870all H ; R oAgua Salud, Panama Canal: USNM 50543234, 50543536both H , 505437, 505438H , 50543940, 505441H , 505442,50544345al l H , 50544649, 505450H , 505451, 505452H ,505453, 50545455both H , 505456; R o Gigantito, PanamaCanal: USNM 50551520, 50552122both H , 505523,505524H , 50552540; R o Mendoza, Panama Canal: USNM50562842, 505643H , 50564448, 505649H ; Tigre Island,Panama Canal: USNM 505484500, 50550105all H ,50550610, 505511H , 50551214; Quipo, on R o Ciri, W side L. Gatu n: AMNH 42919; Darie n: 0.5 hr below junctionof R o Jaque and R o Imamado: USNM 161216; PirriRange, near head of R o Limo n: USNM 50151; Chalichi-mans Creek: AMNH 4291617; Camp Creek: AMNH42920; R o Chucunague [Chucunaque]: AMNH 3790203,49214; R o Chucunague [Chucunaque], 3 mi W of CampTownsend: AMNH 10255758; Yavisa, backyard junkpiles:CM 7403031; Yavisa, trail along R o Chucunague [Chucu-

naque]: CM 74038; 711 km SW El Real between R oPresencia and R o Morgentese, 100350 m: UMMZ15580203; R o Sucubti: AMNH 4290914; Tapia: AMNH25021, 25023, 2502526; Panama : Metropolitan NationalPark, Panama City (8u 589600N, 79u 329460 W), 45 m: SMF85307* H , 85308* , 8530910both * H , 8531119all * ; PanamaProv.: AMNH 71727, 71729, 7600103, 89883; CanalZone: AMNH 67078, 6708182, 71716, 7173033, 75990H ,7599192, 8988082, 8560507, 10743264, 107465H ,107466, USNM 54325; Las Cascadas, Canal Zone: MCZ19414* , 19416* , 175185* , 175187* , 17518990both * ,175194* , 17519697both * , 175200* , 175203* , 175207* ,175209* , 17521316all * , 175223* , 175225* , 175233* ; CanalZone, Camp Mary Caretta (5 Camp Santa Margarita):USNM 2516263; Canal Zone, Gamboa: USNM 193351;Canal Zone, R o Frijoles, 3 mi N Gamboa: UF 124417;Gamboa, at confluence of Panama Canal and Chagres River(9u 069540N, 79u 419420 W): USNM 29780709; Gamboa:SMF 84954, 8308485; Canal Zone, R o Medio: USNM102725; Lion Hill: USNM 54172; Old Panama: USNM50129; Panama, Cabima (Pacific slope): USNM 4850001;Panama, Cocoli: USNM 193365, 193371, 52337778;Puerta Obaldia, Quebrada Repressa: USNM 150127; Venado Beach: USNM 193359; Punta de Pena: USNM38712; Toro Point: USNM 53725; Trinidad River: USNM6399294, 63997; Viento Fr o: USNM 48597; La Joya,Pacific side: ANSP 25136* ; R o Tatare, Pacific side: ANSP25137; Gatu n: ANSP 1952022all * , 19523, 2486365all * ,2486667; Canal Zone, Fort Gulick, Atlantic side: ANSP25104* ; Canal Zone, Empire: ANSP 19545; Chico: CM6859; Barro Colorado Island: AMNH 75986, ANSP 2448792all * , 24493* H , 24494500, 24559, CM 7664, 7666, 7669,7671, 767375, 7681, 768693, 7699, 7705, 770710, 771219, 7725; Barro Colorado Island, Wheeler trail: UMMZ63688; Lutz Creek below Donats bridge: CM 7659; R oPequeni, head of Madden Lake: ANSP 21694; Juan Mina,Madden Lake watershed: CM 74047; near Fort ClaytonReservation: UIMNH 42184; Cerro Campana: AMNH75999; Cerro Campana, 800900 m: AMNH 10666; Altosde Maje : AMNH 109623H , 10962435; Serran a de Maje ,proximities of Unio n Saldan a, R o Chima n (8u 51959.00N,

78u

359

13.60

W), 470 m: MHCH 1146; Serran a de Maje , R oAmbroya: MHCH 1082H , 1086, 1090; San Bla s: Armila:USNM 150099, 150100H , 150101108; Armila, Quebrada Venado: USNM 15011013; Nusagandi, near field station


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(9u 20.509N, 78u 59.649 W), 300360 m: SMF 80717H , 80718,80719H , 80720* H ; km14.6 onEl Llano Cart road, 370 m:AMNH 110572; km 12.8 on El Llano Cart road, 290 m:AMNH 11057374.

Anolis cryptolimifrons.Costa Rica: Limo n: SE sideCerro Nimaso: UCR 8477* H ; Panama: Bocas del Toro:Cerro Brujo (9u 11916.40N, 82u 11925.40 W), 10 m: SMF85230* H , 8523135all * , 8523637both * H , 8523841all * ,8524243both * H , 85244* ; vicinity of Almirante: ANSP3404750all * H , 34051* , 34052H , 3405354, USNM27906271, 27913033; R o Changuinola, near QuebradaEl Guabo, 16 km airline W Almirante, 200250 m:AMNH 119043H , 11904449; Cayo Agua, Punta Norte:USNM 150005, 15000709; Cayo Agua, near PuntaLimo n: USNM 33869092; Isla Bastimentos, Old Point:USNM 29788897; Isla Bastimentos: SMF 85229,85245* H ; Isla Colo n, ca. 0.8 mi N of Bocas del Toro(town): USNM 33821416; Isla Colo n, just N of Bocas delToro (town), along beach at fairgrounds, E side of

isthmus: USNM 346901; Isla Colo n, La Gruta: USNM31376778, 313794, 338217, 338218H , 33821922,338223H , 33822426, 33822729al l H , 33823032; IslaCristo bal, Bocatorito camp: USNM 348191H , 34819294,348195H , 348196201, 348202H , 34820305; NW side of Isla Cristo bal: USNM 348206H , 34820710; Isla Pastores,Ford Point: USNM 31384748; Isla Popa, 1 km SE of Deer Island channel: USNM 29812135; Isla Popa, southend of, 1 km E of Sumwood Channel: USNM 31921325,34726063, 34726465both H , 34726671, 34727275all H ,34727677, 347278H , 347279, 347280H , 34728183; IslaPopa (9u 139140N, 82u 089280 W), 10 m: SMF 8524748both * H , 85249* , 85250* H ; Isla Popa, NNE beach(09u 13924.40N, 82u 06936.60 W), 1020 m: SMF 85399H ,

85400; Laguna de Tierra Oscura, 3.7 km S of Tigre Key:USNM 31383851, 34846770all H , 34847180; Long Bay Point and Flat Rock Point, between, on E side of island,ca. 100 yds from beach: USNM 29781617; midpoint on W side of Cayo Carenero: USNM 347938; N end of CayoRoldan: USNM 348043; Isla Solarte, 10 m: SMF85251* H , 8525253both * ; USNM 338552, 33855355all H ,33855657, 338558H , 33855961, 33856263both H ,338564, 338565H , 33856668.

Anolis limifrons.Costa Rica: Costa Rica: USNM38334, 7040610, 81198; Parismina nivel del mar: USNM7544446; Colombiana: USNM 6734748; Alajuela: R oFr o: USNM 19514; Pizote: UCR 998889both * H ,10646* H ; Pilo n, Bijagua: UCR 10504* H ; Laguna LagartoLodge, Boca Tapada: UCR 12609* H ; R o Tapezco: UCR16515* H ; Cartago: 10.0 km NE Turrialba on E bank of R o Reventazo n R1287375; 2.0 km W Pavones deTurrialba R1287879; 3.0 km NE Pavones de Turrialbaat R o Chitar a R1286972, R1287677; Tapanti: ZFMK48716; Turrialba Prov.: AMNH 6970710; Turrialba:SMF 77206, USNM 133180, 192586, 52337576; Esta-cio n Biolo gica Copal, Tausito, Pejibaye: UCR 16127* ;1 Km E La Pastora (9u 58906.10N, 83u 44918.90 W), 1500 m:SMF 86924; Heredia: Rara Avis, Catarata (10u 16.929N,84u 02.749 W), 700 m: SMF 81814, 81815H ; Puerto Viejo:ZFMK 4872337, USNM 245041; La Selva BiologicalStation, 2.6 km SE of Puerto Viejo de Sarapiqu : USNM

50567790, 505691H

, 50569295, 505696H

, 505697,505698H , 505699701; Finca Santiago (near La Selva):SMF 78433* ; Limo n: Limo n Prov.: AMNH 8917173,8917576, 95095, 9967176, 14961120; ca 5/4 mi S

mouth of R o Tortuguero, ca. 50 mi NW Limo n: AMNH89174H ; ca. 5 mi N Limo n: AMNH 89177H ; Zent[10u 01960 N, 83u 16960 W, 31 m]: USNM 137767;Approximately 17.0 km WSW Puerto Limo n betweenR o Blanco and R o Toro R1288288; Siquirres largestream outside of town R1288081; Motel Matama,3.5 km N Limo n: AMNH 13860405; 4 mi SW LaFortuna: IRSNB 11684; Atalanta Farm, Estrella Valle:ANSP 21465; Puerto Limo n: ANSP 19570, 19571H ,1957278, IRSNB 13804, ZSM 85/1998, 86/1998; LaCastilla, lower R o Reventazo n: ANSP 2371037, 2450104, 34747; Tortuguero: UF 13578384; Tortuguero, just Nof Caribbean Conservation Commission Camp: USNM244861; 2.4 km E Siquirres, along R o Pacuare: CM8956667; Rte 32, 69 km E R o Hondo, 2 km N on dirtroad: ANSP 32372, 32374; 23 km (air) NW Bribri at R oCarbo n along road to Uatsi: ANSP 32559; RB Hitoy Cerere: SMF 86925* H , 86926* , 8692728both * H ; Estacio nBiolo gica Tierra Media, Matina: UCR 12399* H ; Quebrada

Uatsi: UCR 13031* H

; Sendero San Mateo, Cerro Uatsi:UCR 13195* H ; Finca Bryan Kubicki, Guayaca n: UCR16914* H ; Puntarenas: Puntarenas Prov.: AMNH 16357;7 mi E Golfito: LSUMZ 30260; Pen nsula de Osa, GolfoDulce, Puerto Jime nez, jardin at Jime nez Yacht Club:SMF 8151215; EB San Gerardo, Monteverde: UCR13652* H ; Hotel Sunset, 1 km N Santa Elena, Monteverderegion (10u 19936.90N, 84u 49924.10 W), 14501475 m:SMF 8554953all * , 85554* H ; San Jose : San Jose : ANSP7804, USNM 8090205; San Jose , grounds of Hotel Irazu :UMMZ 143761; 6.0 km N San Isidro de Perez Zeledo n:UTA R12868; Talamanca: USNM 75956; Moravia deChirripo: UMMZ 128952; Near San Isidro (9u 24929.10N,83u 44906.60 W), 880 m: SMF 8693335; Road fromGeneral Viejo to Santa Elena (9

u

20906.50N,83u 39911.40 W), 650 m: SMF 86929;Honduras: Colo n:Quebrada Mach n (15u 199100N, 85u 179300 W), 540 m:USNM 53649091, 54102629; R. B. R o Pla tano, ElOcotillal, Cabeceras de R o Pla tano (15u 40.39N,85u 17.19 W) 370410 m: SMF 86215H , 86216, 86217H ;Gracias a Dios: Mocoro n R4617172; confluence of R o Wampu and Quebrada Waskista (15u 009N, 84u 599 W),85 m: USNM 33018384; confluence of R o Wampu andR o Patuca (14u 589N, 84u 599 W), 60 m: USNM 33018182; Quebrada Waskista, 85 m: SMF 8070809both * ; R. B.R o Pla tano, Raudal Kiplatara, (15u 59.99N, 84u 94.89 W),50255 m: SMF 86172H , 8617374, 86175H , 8617682,8618384both H , 86186; R. B. R o Pla tano, R o Cuyamel(15u 58.29N, 84u 99.39 W), 115345 m: SMF 86188,86189H , 86190, 86191H , 8619296, 86198H , 8620405;R. B. R o Pla tano, Pomokir (15u 49.39N, 84u 94.89 W), 150240 m: SMF 86207, 8620809both H , 86211, 8621314both H ;R. B. R o Pla tano, Crique Unawas (15u 12.79N, 84u 92.39 W),180305 m: SMF 8622021, 862223both H , 8622428,86229H ; Quebrada Waskista-R o Wampu confluence(15u 009N, 84u 599 W), 85 m: SMF 86887; Cabeceras delR o Rus Rus, 190 m: SMF 86888; Olancho: confluence of R o Wampu and Quebrada Siksatara (15u 039N, 85u 029 W),95 m: USNM 330180; confluence of R o Aner and R o Wampu (15u 049N, 85u 069 W), 110 m: SMF 80704, USNM33017677; confluence of R o Wampu and R o Sausa

(15u

049

N, 85u

069

W), 100 m: SMF 80705, 8070607both *

,USNM 33017879; confluence of R o Yanguay and R o Wampu (15u 039N, 85u 089 W), 110 m: USNM 330175;Parque Nacional Patuca, Matamoros (14u 409210N,


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85u 239110 W), 150 m: SMF 80710, 80712* ; Parque NacionalPatuca, Quebrada El Gua simo (14u 349380N, 85u 179540 W),140 m: SMF 80713* , 8071415, 80716H ; Parque NacionalPatuca, Caobitas (14u 399220N, 85u 179430 W), 100 m: SMF80711; Quebrada Siksatara-R o Wampu confluence(15u 049N, 85u 029 W), 95 m: SMF 8688990; R o Yanguay-R o Wampu confluence (15u 039N, 85u 089 W), 110 m: SMF8689193; R o Aner-R o Wampu confluence (15u 039N,85u 079 W), 110 m: SMF 8689496; R o Sausa-R o Wampu confluence (15u 049N, 85u 069 W), 100 m: SMF 86897900,86901H , 8690203; Yapuwa s (14u 589N, 85u 009 W), 60 m:SMF 8690406; Between R o Sausa and R o Wampu ,100 m: SMF 86907, 86908; Quebrada de Las Mar as(15u 189N, 85u 219 W), 660 m: SMF 86909H ; Matamoros(14u 409N, 85u 239 W), 150 m: SMF 8691014all H ; QuebradaEl Mono (14u 399N, 85u 209 W), 100 m: SMF 86915;Quebrada El Gua simo (15u 059N,86u 259 W), 140 m: SMF86916; Nicaragua: no specific locality: USNM 13739,15212; Palvon (5 El Polvo n?): USNM 120758; Atla ntico

Norte: Eden Mine: ANSP 21138*

; Great Falls, Pispis:ANSP 21124* ; Parque Nacional Saslaya, Estacio n BiologicaSalto Labu (13u 399510N, 85u 00955.50 W), 260 m: SMF8206869both * ; Parque Nacional Saslaya, between Estacio nBiologica Salto Labu and Campamento El Revenido, 400500 m: SMF82236; ParqueNacional Saslaya, CampamentoI (13u 439110N, 85u 029200 W), 720 m: SMF 82070H ; ParqueNacional Saslaya, Campamento El Carao (13u 42.799N;84u 58.669 W), 400 m: SMF 79611; Parque Nacional Saslaya,Campamento Las Pavas (13u 44.59N; 85u 01.59 W), 780 m:SMF 79379H ; Parque Nacional Saslaya, trail from Campa-mento Las Pavas to Campamento El Carao: SMF 82844;Parque Nacional Saslaya, trail from Campamento Las Pavas(13u 44.59N; 85u 01.59 W) to Campamento Los Monos(13

u

45.19N; 85u

02.29 W), 810 m: SMF 79382* H

; ParqueNacional Saslaya, Campamento Los Monos (13u 45.19N;85u 02.29 W), 800820 m: SMF 79161, 79591* , 79607,79898* ; Parque Nacional Saslaya, Campamento Las Ranas,920 m: SMF 82071* ; Cerro El Toro (13u 42930.60N,85u 02917.70 W), 830 m: SMF 83209; Rancho Alegre(13u 39947.50N, 85u 01938.90 W), 285 m: SMF 84737* , JS052* ; Finca URACCAN (13u 43944.10N, 84u 53914.50 W),145 m: SMF 84738* , JS 074* ; 4 km E Bonanza, 200 m: KU101397; Bonanza, 260 m: KU 84867, 8565657, 101398400; Alamikamba (13u 30.089N, 84u 13.649 W): SMF 7755255; Moss, 2 km S Finca Luciana, along R o Wawa: JS 1089,109091both H , 1113H , 1130; Krin Krin, 5 km W confluenceR o Waspuk with R o Coco: JS 1193; Atla ntico Sur: R oEscondido, 45 mi. from Bluefields: USNM 1973334; R oEscondido, 50 mi from Bluefields: USNM 19877; Cara deMono, 50 m: KU112991; ElRecreo, S sideR o Mico, 25 m:KU 10186567, 11298890; vicinity of Providencia(11u 31.799N, 84u 22.929 W), 100 m: SMF 77556; Boaco:Finca Santa Helena, Masigue, 30 km NE Camoapa, 600 m:SMF 84733* ; Jinotega: Finca Berl n (13u 32.269N,85u 41.509 W), 1015 m: SMF 78980* H ; Cerro Kilambe , LaCueva (13u 36911.10N, 85u 42952.60 W), 1025 m: JS 161* ;Cerro Kilambe , El Diamante (13u 36951.30N, 85u 44920.20 W,1090 m: SMF 84739* ; Cerro Kilambe (13u 37906.20N,85u 43919.50 W, 1015 m: SMF 84740* H ; Reserva BiosferaBosawas, ca. 0.5 km SE Pueblo Wiso (13u 59.609N,85u 19.600 W), 200 m: SMF 7853435both * H ; Reserva Bios-

fera Bosawas, Ayapal (13u 46.619N, 85u 24.149 W), 195 m:SMF 78404* ; Cordillera Isabelia (13u 19.999N,85u 41.529 W), 500 m: SMF 78314H , 78315, 78316H ; FincaSanta Enriqueta, 5 km E San Jose De Las Latas, 1300 m:SMF 84732* H , JS 061* ; Rait , along R o Coco: JS 919921;Matagalpa: Selva Negra (12u 59.969N, 85u 54.559 W): SMF7720203both * , 7720405, 77334, 77551, 78201, 78202H ,7820304, 78205H ,78206, 78281, 78312, 79814* H ; 12 kmNE Matagalpa, 1100 m: KU 19507778; Finca Tepeyac,10.5 km N and 9 km E Matagalpa, 960 m: KU 8565155,85675; road to Puerto Cabezas (13u 17.119N, 85u 42.949 W),1000 m: SMF 78313; Cerro Musu n, FUNDENIC(12u 57918.80N, 85u 13951.20 W), 630 m: JS 737H , 740,753H ; R o San Juan: Machuca: ANSP 7803* ; at Isla deDiamante on R o San Juan: OMNH 3593233; R o SanJuan: USNM 2498182; SanJuan del Norte: USNM 1950607; R o San Juan, Boca de San Carlos (10u 47.269N,84u 11.709 W), 20 m: SMF 79815* , 79816H ; Boca de SanCarlos (10u 47925.70N, 84u 11937.70 W), 40 m: JS 528; R oSan Juan, Bartola (10

u

58.379N, 84u

20.359 W), 30 m: SMF79817* H , 79818, 8092439, 80965; Boca de Bartola(10u 58918.30N, 84u 20923.10 W), 25 m: JS 454H , 455; conflu-ence of R o San Juan and R o Sarapiqu : SMF 83206H ,83207; Chingo Petaca (10u 44950.90N, 83u 50926.30 W), 40 m:SMF 83208H ; Dos Bocas de Bartola, El Almendro(10u 59943.90N, 84u 16937.50 W), 75 m: JS 402H ; Dos Bocasde R o Indio (11u 02954.80N, 83u 52948.40 W), 25 m: JS 542H ,543, 622H ; R o Fr o, FUNDEVERDE (11 u 04937.00N,84u 44955.70 W), 45 m: JS 327H , 328, 330; R o Fr o,FUNDEVERDE (11 u 03941.80N, 84u 44927.00 W), 50 m: JS355; along R o Papaturro, Refugio de Vida Silvestre LosGuatuzos: JS 953H , 954, 958H ; Islas de Solentiname: KU17404850, SMF 77210, 77660, 7832430; Islas deSolentiname, Isla Mancarro n, Hotel Mancarro n: SMF82197H ; Islas de Solentiname, Isla Mancarro n: SMF 82198,82199* ; Islas de Solentiname, Isla Mancarro n, 1 km NHotel Mancarro n: SMF 82200* H ; Panama: Bocas delToro: 7.5 km airline WSW Chiriqu Grande, 10 m: AMNH123265, 123266H , 12326768, 123269H ; Rambala, nearChiriqu Grande (08u 559270N, 82u 119030 W), 20 m: SMF85254H , 85255; Isla Escudo de Veraguas, West Point:USNM 34748389, 347490H , 347491; Celentine(8u 479090N, 82u 119170 W), 610 m: SMF 85228* H ; QuebradaLa Gloria, near Miramar (8u 59908.00N, 82u 13956.40 W),20 m: SMF 85246* H ; Bocadel R o Krikamola (8u 58938.60N,81u 55901.70 W), 7 m: MHCH 539H ; Chiriqu : Chiriqu :

ZMB 7785, 7827; ca. 6 mi below El Hato, alongHwy, 4000 ft: FMNH 60275; Bugabita, 100200 m:NMW 20673 (13); Boquete: ZSM 63/1989/3; David,UNACHI, Jard n Bota nico, near River: SMF 85260* ;Los Algarrobos (8u 29939.10N, 82u 26901.30 W), 110 m:SMF 85223* ; Los Algarrobos (8u 299250N, 82u 269100 W),130 m: SMF 85256* ; Los Algarrobos (8u 299470N,82u 269000 W), 130 m: SMF 8525759all * ; Meseta deChorcha (8u 249380N, 82u 139100 W), 190 m: SMF 85227* ;Meseta de Chorcha (8u 249500N,82u 139060 W), 240270 m:SMF 8522426all * ; Cocle : N El Valle de Anto n, CerroGaita l (8u 37.679N, 80u 6.609 W), 750 m: SMF 80779; Veraguas: 56 mi (via road) NW Santa Fe: AMNH120003H .


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i0018-0831-64-1-92