Anolis apletophallus und cryptolimifrons köhler&sunyer 2008

download Anolis apletophallus und cryptolimifrons köhler&sunyer 2008

If you can't read please download the document

Transcript of Anolis apletophallus und cryptolimifrons köhler&sunyer 2008

  • 1. Herpetologica, 64(1), 2008, 92108E 2008 by The Herpetologists League, Inc.TWO NEW SPECIES OF ANOLES FORMERLY REFERRED TO ASANOLIS LIMIFRONS (SQUAMATA: POLYCHROTIDAE) GUNTHER KOHLER1,3 ANDJAVIER SUNYER1,21Forschungsinstitut und Naturmuseum Senckenberg, Senckenberganlage 25, 60325 Frankfurt a.M., Germany2 n, Gabinete de Ecologa y Medio Ambiente, Departamento de Biologa, UNAN-Leo Nicaragua ABSTRACT: We describe two new species of anoles (genus Anolis) from Panama formerly referred to as Anolis (or Norops) limifrons. Both new species differ from A. limifrons by having a large bilobed hemipenis (small and unilobed in A. limifrons). The new species differ from each other in male dewlap size and coloration. We provide an identification key and standardized descriptions of A. limifrons and the two new species described herein. RESUMEN: Describimos dos especies nuevas de anoli (genero Anolis) de Panama anteriormente referidas como Anolis (o Norops) limifrons. Ambas nuevas especies difieren de A. limifrons en tener un hemipene grande y bilobulado (pequeno y unilobulado en A. limifrons). Las dos especies nuevas difieren entre s en el tamano y coloracion de la papera gular de los machos adultos. Se proporciona una clave dicotomica de identificacion y descripciones estandar de A. limifrons y de las otras dos nuevas especies.Key words: Anolis; New species; Panama; Polychrotidae; Reptilia; Squamata ANOLES are a diverse and taxonomically an adult male specimen (now ANSP 7890).poorly understood group of lizards that are According to Savage (1973:36), the holotypedistributed widely across the tropical andof A. bransfordii was collected by Bransfordsubtropical portions of North, Central andat Machuco (5 Machuca) on the Ro San South America (Kohler, 2003; Savage, 2002).Juan, Departamento Ro San Juan, Nicara-Several new species have been described ingua. In 1882, Thominot described Anolisrecent years indicating that more field workrivieri based on a juvenile specimen (nowand study of museum material is needed to MNHN 1884.221) from Panama. Boulengerdocument the real diversity of anoles. The(1885) described Anolis godmani based oncomparative study of hemipenial variation infour specimens from Guatemala and threeanoles revealed substantial cryptic diversityspecimens from Irazu, Costa Rica. Stuart (Kohler and Kreutz, 1999; Kohler et al., 2003). (1955:30) presumed that in Anolis godmani In 1862, Cope described the new speciesBoulenger [there] may have been a mixupAnolis (Dracontura) limifrons based on twoin locality data in the Godman-Salvin collec-syntypes (now ANSP 790001) from Vera-tions, one of the cotypes having been listed asgua. According to Savage (1970:279), todayof Guatemala whereas it probably came fromthe old Veragua comprises the Provincias of Costa Rica, and he also stated that theVeraguas, Chiriqu and Bocas del Toro.material [including the holotype of A. god-Barbour (1934:139) described the type localitymani] may have been received by the Britishof A. limifrons as Cucuyos, Veragua Prov.,Museum somewhat after the main bulk of thePanama [an abandoned mine on the Ro earlier parts of the collection had been turnedSantiago]. In 1871, Cope named Anolis over to the Museum. In 1956, Taylortrochilus based on an adult male specimen described the new species Anolis biscutiger(now ANSP 7804) from San Jose, Costa based on an adult male (KU 40771) fromRica. Peters (1873) added another nominal Golfito, Puntarenas Province, Costa Rica.species, Anolis pulchripes, based on a speci- Dunn (1930) regarded rivieri, trochilus andmen (now ZMB 7827) from Chiriqu. A yearbransfordii as synonyms of limifrons. Barbourlater, Cope (1874) described the new species(1934:140) stated that Dunn has seen theAnolis bransfordii from Nicaragua based ontypes of limifrons, rivieri, pulchripes androdriguezii and declares them all the same3CORRESPONDENCE: e-mail, [email protected] species. The names trochilus, pulchripes, 92

2. March 2008] HERPETOLOGICA93bransfordii and rivieri have remained in the in order to document any variation. Wesynonymy of A. limifrons Cope whereas god- measured scale sizes using the ocular micro-mani has been retained as a valid species untilmeter of a stereo microscope (Leica MZ 12)recently (Peters and Donoso-Barros, 1970;and rounded the values to the nearestSavage and Villa, 1986; Taylor, 1956; Villa et 0.01 mm. For all other measurements we usedal., 1988). Savage (2002) and Kohler (2003)precision calipers and rounded the values torecognized a northern species, rodriguezii,the nearest 0.1 mm. We measured head lengthand a southern one, limifrons, with biscutiger from tip of snout to the anterior margin of theand godmani as synonyms of limifrons.ear opening. We measured snout length from Here we report the results of our study oftip of snout to the anterior border of the orbit.the variation in hemipenial and scalationWe determined head width as the distancemorphology as well as morphometrics of the between rictuses. We counted dorsal andsmall anoles occurring from eastern Honduras ventral scales at midbody along the eastern Panama commonly referred to asWe measured tail height and width at the pointAnolis (or Norops) limifrons.reached by the heel of the extended hind leg. We counted subdigital lamellae on phalanges ii MATERIALS AND METHODS to iv of the 4th toe. We considered the scale directly anterior to the circumnasal to be aFor this study, we examined 1428 specimens prenasal. The capitalized colors and colorof Anolis limifrons. We provide a list of the codes (the latter in parentheses) are those ofcomparative specimens examined in the Ap- Smithe (19751981). To measure dewlap area,pendix. Abbreviations for museum collections we took photographs of males with theirfollow those of Leviton et al., (1985) except dewlaps artificially extended using small fore- MHCH (Museo Herpetologico de Chiriqu, ceps. The head portion was magnified and David, Panama), and JS field numbers, which printed, then superimposed on millimetricrefer to specimens that will be deposited in the paper, and the total number of 1-mm squaresMuseo de Ciencias Naturales de la Universidad contained in the extended dewlap counted. ACentroamericana (UCA), Managua, Nicara- straight line was drawn between both thegua. In the course of this study, G.K. had the anterior and posterior insertions of the dewlap.privilege of examining all extant primary types We also determined the HL on the printout.of nominal species regarded as synonyms of We used the following equation to convert the Anolis limifrons by previous authors (Kohler, magnified dewlap area to the real size: X 52003; Peters and Donoso-Barros, 1970; Savage, [(!Y / A) ? B]2; where: X is the real area of the2002). For the synonymy lists, we included only dewlap in mm2; Y is the total area (mm2) of thethose works that cite actual specimens. No- dewlap at a magnified scale; A is the HLmenclature of scale characters follows that of measure (mm) of the anole at a magnified Kohler (2003). Terminology for hemipenial scale; and B is the HL measure (mm) of themorphology follows that of Myers et al. (1993) anole at the real size. We executed statisticaland Savage (1997). We everted the hemipenes techniques using the computer program Sta-in the field after euthanasia and before tistica version 6.1. Juveniles were not includedpreserving the lizards in 70% ethanol by in the statistical analyses. We used discrimantmanually applying pressure to the ventral base function analysis to evaluate the pheneticof tail. Once the hemipenes had been everted distinctness of a priori groups. Discriminantto some extent, we injected 70% ethanol in the scores (DC) were calculated by multiplyingarea of the hemipenial pockets, usually selected variables by their associated canonicalresulting in complete eversion of both hemi- coefficients. Each specimen was then plottedpenes. With the needle still attached to the along the axae providing maximal separation oflizards tail, we placed the lizard for 12 min the a priori groups.into the preservative to allow fixation of theeverted hemipenis. We did not tie off thehemipenes at the base because this might causeRESULTSdamage to these fragile organs. We intended to Two distinctly different hemipenial mor-evert the hemipenes in all available adult males photypes are evident in the specimens we 3. 94 HERPETOLOGICA [Vol. 64, No. 1examined. In Type A (n 5 188 adult males(Fig. 1) that correctly classified 72.7% of thewith everted hemipenes), the hemipenis is a specimens. The first function is DS 5 0.471297relatively large bilobed organ; both the(number of medial ventral scales in one headtruncus and the lobes have a strongly calycu- length) +0.673627 (number of medial dorsallate surface. The sulcus spermaticus bifurcates scales in one head length) 20.392336 (sub-at the base of the apex and the branchesdigital lamellae) +0.176923 (total number ofcontinue to the tip of the lobes. In Type B (nloreal scales) +0.286056 (scales between su-5 85 adult males with everted hemipenes), praorbital semicircles) 20.651674 (scales be-the hemipenis is much smaller relative to bodytween posterior canthals) +0.025958 (scalessize as compared to the Type A hemipenis. around midbody). The second function is DSAlso, it is unilobed and the truncus and lobes5 0.284551 (number of medial ventral scales inare not calyculate. The sulcus spermaticusone head length) 20.173192 (number ofopens at the base of the apex. While these twomedial dorsal scales in one head length)hemipenial types show a broadly sympatric 20.610593 (subdigital lamellae) 20.171498geographical distribution pattern in western(total number of loreal scales) 20.293526Panama, variation in hemipenial morphology(scales between supraorbital semicircles)within these discrete types is negligible both+0.054329 (scales between posterior canthals)within populations and in a geographical20.603442 (scales around midbody).context. Within Type A, two distinct types Based on our data, we recognize threecan be distinguished in respect of the relative species of this complex: Species A: Hemipenissize and the coloration of the male dewlap. unilobed; male dewlap small (smaller thanMales with bilobed hemipenes from the 100 mm2), dull white with a small basalProvince of Bocas del Toro, Panama, and orange blotch; distributed from eastern Hon-adjacent southeastern Costa Rica have a small duras to central Panama west of the Canaldewlap (smaller than 100 mm2) that is dullZone (Fig. 2). Species B: Hemipenis bilobed;white with a small basal orange blotch. Males male dewlap small (smaller than 100 mm2),with bilobed hemipenes from central and dull white with a small basal orange blotch;eastern Panama have a large dewlap (largerdistributed in the western and central portionsthan 150 mm2) that is almost uniformlyof the Province Bocas del Toro, Panama, andorange. All Type B males have a small dewlapadjacent southeastern Costa Rica (Fig. 2).(smaller than 100 mm2) that is dull white withSpecies C: Hemipenis bilobed; male dewlapa small basal orange blotch.large (larger than 150 mm2), almost uniformly In external morphology, there is great orange; distributed in central and easternoverlap in the ranges of all examined charac- Panama; expected in adjacent northwesternters of scalation and morphometrics (Table 1).Colombia (Fig. 2). In western Panama (Prov-We conducted a discriminant function analy- inces of Chiriqu, Bocas del Toro andsis (DFA) based on seven pholidotic charac- Veraguas), the bilobed form (our Species B)ters (number of medial ventral scales in oneis restricted to the Caribbean versant; whereashead length; number of medial dorsal scales inthe unilobed form (our Species A) occurs onone head length; subdigital lamellae; total both versants. Although no cases of actualnumber of loreal scales; scales between syntopy of these two forms have beensupraorbital semicircles; scales between pos- documented, they show a broadly sympatricterior canthals; scales around midbody) and distribution pattern on the Caribbean versantdid a priori assignments to groups based on of western Panama.our hemipenis and male dewlap findings Unfortunately, none of the type specimens(Group 1 5 Type B hemipenis, male dewlapmentioned in the introduction are males withsmall, dull white with a small basal orange their hemipenes everted. Also, data on dewlapblotch; Group 2 5 Type A hemipenis, malecoloration in life is not available for these typedewlap small, dull white with a small basal specimens. However, the type locality data oforange blotch; Group 3 5 Type A hemipenis,most of these nominal species allow for amale dewlap large, more or less uniformly reliable allocation to one of the three speciesorange). This DFA yielded a scatter diagram that we distinguish. Thus, on geographic 4. TABLE 1.Selected measurements, proportions and scale characters of Anolis limifrons, A. apletophallus, and A. cryptolimifrons (specimens with * listed in Appendix). Range is March 2008]followed by mean value and one standard deviation in parentheses. Morphomentric data were only taken from adults. A. limifronsA. apletophallus A. cryptolimifrons = 50 = 25= 20 R 21R 24R 10Maximum SVL =41.547.0 45.0R43.546.5 42.0Tail length / SVL =1.532.52 (2.21 6 0.19) 1.522.29 (2.03 6 0.17)1.832.19 (2.07 6 0.10)R1.602.30 (2.05 6 0.22) 1.822.32 (2.06 6 0.13)1.872.08 (1.98 6 0.08)Tail diameter vertical / horizontal =1.001.30 (1.16 6 0.07) 1.031.27 (1.14 6 0.08)1.001.25 (1.15 6 0.06)R1.001.27 (1.14 6 0.07) 1.001.25 (1.11 6 0.07)1.001.17 (1.21 6 0.06)Head length / SVL =0.250.28 (0.27 6 0.01) 0.250.29 (0.27 6 0.01)0.250.29 (0.27 6 0.01)R0.240.28 (0.26 6 0.01) 0.250.29 (0.27 6 0.01)0.260.28 (0.27 6 0.01)Head length / head width=1.441.80 (1.67 6 0.07) 1.541.84 (1.67 6 0.06)1.591.77 (1.67 6 0.06)R1.571.77 (1.67 6 0.06) 1.561.70 (1.65 6 0.04)1.621.75 (1.66 6 0.04)Interparietal plate / ear =0.795.20 (2.03 6 0.78) 0.714.12 (1.71 6 0.70)0.542.29 (1.47 6 0.46)R1.094.75 (2.19 6 0.98) 1.012.45 (1.69 6 0.44)0.501.68 (1.09 6 0.39)Shank length / SVL=0.260.33 (0.30 6 0.02) 0.290.32 (0.30 6 0.01)0.260.31 (0.29 6 0.02)R0.260.31 (0.28 6 0.01) 0.260.32 (0.30 6 0.02)0.270.29 (0.28 6 0.01)Axillagroin distance / SVL =0.360.47 (0.42 6 0.03) 0.340.46 (0.39 6 0.03)0.390.46 (0.43 6 0.02)R0.410.53 (0.45 6 0.02) 0.350.48 (0.41 6 0.03)0.400.46 (0.43 6 0.02) HERPETOLOGICASubdigital lamellae of 4th toe 2027 (23.33 6 1.61) 2126(23.59 6 1.12)2329(25.03 6 1.47)Number of scales between supraorbital semicircles04 (1.78 6 0.84)13(2.04 6 0.50)13 (2.20 6 0.61)Number of scales between interparietal plate and supraorbital semicircles15 (2.68 6 0.86)14(2.72 6 0.68)24 (3.27 6 0.64)Number of scales between subocular scales and supralabial scales 0 00Number of supralabial scales to level below center of eye58 (6.48 6 0.69)68(6.76 6 0.56)57 (6.73 6 0.52)Number of infralabials to level below center of eye58 (6.30 6 0.70)58(6.37 6 0.57)68 (6.62 6 0.56)Total number of loreals2468 (41.13 6 9.56) 2574(45.00 6 8.78) 3669 (49.07 6 7.67)Number of horizontal loreal scale rows 58 (6.10 6 0.79)58(6.38 6 0.73)68 (6.70 6 0.70)Number of postrostrals 59 (6.86 6 0.82)68(7.23 6 0.59)69 (7.17 6 0.53)Number of postmentals48 (6.45 6 0.91)69(7.28 6 0.88)68 (7.03 6 0.87)Number of scales between nasals 711 (8.91 6 0.86) 811(9.69 6 0.66) 912 (9.77 6 0.77)Number of scales between 2nd canthals 715 (10.59 6 1.69)913(10.61 6 1.06)916 (12.00 6 1.55)Number of scales between posterior canthals1018 (13.77 6 2.10) 1018(13.23 6 1.86) 1219 (15.21 6 1.76)Number of medial dorsal scales in one head length3870 (54.26 6 5.76) 4274(60.65 6 7.10) 4670 (58.87 6 5.35)Number of medial ventral scales in one head length 2658 (40.62 6 6.15) 3660(45.78 6 5.44) 3450 (41.31 6 4.01) 95 5. 96HERPETOLOGICA [Vol. 64, No. 1 (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 et al., (1998); Poe (2004; in part).Anolis limifrons limifrons: Etheridge (1959; in part).Anolis pulchripes Peters, 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 McCranieFIG. 1.Discriminant function analysis of the Central (1994); Kohler (1999); Kohler (2001); Koh- American anoles formerly referred to as Anolis limifrons. ler et al., (2001); Kohler and McCranieSee text for details.(2001). Diagnosis.Amedium-sized speciesreasons the following taxa (respective type (snoutvent length [SVL] in largest specimenlocalities in parentheses) are clearly referable43.5 mm) of the genus Anolis (sensu Poe,to our Species A: Anolis limifrons Cope 2004) that is most similar in external mor-(Cucuyos, Veragua Province, Panama); A. phology to a cluster of Central American,trochilus Cope (San Jose Costa Rica); A.species that are long-legged (longest toe of pulchripes Peters (Chiriqu, Panama); A.adpressed hindlimb reaches to mid-eye orbransfordii Cope (Nicaragua); A. godmani: beyond), have a single elongated prenasal Boulenger (Irazu, Costa Rica); and A. biscuti-scale, smooth to slightly keeled ventral scales,ger Taylor (Golfito, Puntarenas Province, Costa and slender habitus, often delicate (i.e., AnolisRica). Because A. limifrons Cope is the oldestdollfusianus, A. ocelloscapularis, A. rodrigue-available name for this species, our Species Azii, A. yoroensis, A. zeus). Within this clusterhas to be referred to that name and the other of species, A. limifrons can be readilynominal species remain in the synonymy of A.distinguished by male dewlap coloration (dulllimifrons. The holotype of A. rivieri Thominotwhite with a small basal orange blotch in a juvenile with unspecific locality data limifrons vs. uniformly dull white in A. zeus,(Panama), and its taxonomic identity cannot and almost uniformly orange to orange-yellowbe determined. Therefore, we consider the remaining species. Additionally, A.rivieri Thominot to be a nomen dubium.limifrons differs from the species in thisInterestingly, there is no available scientificcluster by the following characteristics (con-name for either of our Species B and C. Wedition for A. limifrons in parentheses): Anolistherefore describe them as new species below.dollfusianus: ventrals weakly keeled (smooth). Anolis limifrons Cope, 1862Anolis ocelloscapularis: An ocellated shoulderspot present (absent); ventrals weakly keeledAnolis biscutiger Taylor, 1956:81. Holotype (smooth); hemipenis bilobed (unilobed). Ano-KU 40771 from Golfito, Puntarenas Prov- lis rodriguezii: hemipenis bilobed (unilobed).ince, Costa Rica. Anolis yoroensis: ventrals weakly keeledAnolis bransfordii Cope, 1874:67. Holotype(smooth).ANSP 7890 from Nicaragua.Description (Fig. 3).Maximum SVL 41.5Anolis godmani Boulenger, 1885:85. Syntypes mm in males, 43.5 mm in females; ratio tail(BMNH and MCZ) from Guatemala and length/SVL 1.532.52 (2.17 6 0.21); tailIrazu, Costa Rica. Barbour (1834), Taylor slightly compressed in cross section, ratio tail(1956). height/tail width 1.001.30 (1.16 6 0.07); ratioAnolis limifrons Cope, 1862:178. Syntypes axilla to groin distance/SVL 0.360.53 (0.43 6ANSP 790001 from Veragua. Bocourt0.03); ratio head length/SVL 0.240.28 (0.26 6. March 2008] HERPETOLOGICA97FIG. 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 symbolcan represent one or more nearby localities. Areas above 500 and 1000 m are shaded gray. Open symbols: specimenswith no everted hemipenis; Solid symbols: specimens with everted hemipenis.6 0.01); ratio snout length/head length 0.41hind limbs; scales on snout varying from0.52 (0.45 6 0.02); ratio head length/head almost non-keeled to keeled; 59 (6.9 6 0.8)width 1.441.80 (1.67 6 0.06); longest toe ofpostrostrals; 711 (8.9 6 0.9) scales betweenadpressed hind limb reaching to a pointnasals; 1 large elongated prenasal scale inbetween anterior to eye and tip of snout; ratiocontact with both rostral and first supralabial,shank length/SVL 0.260.33 (0.29 6 0.02);occasionally only in contact with rostral; scalesratio shank length/head length 0.971.24 (1.11 in distinct prefrontal depression generally6 0.07); longest finger of extended forelimb slightly tuberculate posteriorly, wrinkled an-reaching to a point between nostrils and tip ofteriorly, some of them keeled; supraorbitalsnout; longest finger of adpressed forelimbsemicircles well developed, separated by 04reaches in between anterior to insertion of(1.8 6 0.8) scales; supraorbital disc composedhind limbs and slightly beyond to insertion of of 514 distinctly enlarged keeled scales; 7. 98HERPETOLOGICA [Vol. 64, No. 1to 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 incontact with supralabials; ear opening medi-um-sized, ratio tympanum height/interparietalscale length 0.581.36 (0.86 6 0.16); mentaldistinctly wider than long, completely dividedmedially, bordered posteriorly by 48 (6.5 60.9) postmentals; 58 (6.3 6 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 toaxilla; dorsum of body with weakly keeledgranular scales (at least anteriorly) withrounded posterior margins, 2 medial rowsslightly enlarged, 3870 (54.3 6 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 (64.6 6 5.7) ventral scalesbetween axilla and groin; 108157 (132.1 610.6) scales around midbody; caudal scalesFIG. 3.Head of Anolis limifrons (SMF 86900). Scalestrongly keeled; caudal middorsal scalesbars equal 1.0 mm.slightly enlarged, without whorls of enlargedscales, although an indistinct division incircumorbital row usually incomplete, there-segments is discernible; a pair of slightlyfore, 03 enlarged supraorbitals in contact enlarged postanal scales usually present; nowith supraorbital semicircles; a single large tube-like axillary pocket present; scales onelongated superciliary; 36 rows of small dorsal surface of forelimb keeled, imbricate;keeled scales extending between enlargeddigital pads dilated; distal phalanx narrowersupraorbitals and superciliaries; a very shallowthan and raised from dilated pad; 2027 (23.3parietal depression present in most speci-6 1.6) lamellae under phalanges iiiv ofmens; interparietal scale well developed, fourth toe; 711 (8.5 6 0.9) scales underusually surrounded by scales of moderate size distal phalanx of fourth toe.anteriorly and by small to moderate size scalesDescription of completely everted hemipe-posteriorly; 15 (2.7 6 0.9) scales present nis.Small unilobed organ; sulcus spermati-between interparietal and supraorbital semi-cus bordered by well developed sulcal lips andcircles; canthal ridge distinct, composed of 6 opens at base of apex; no discernable surface11 (7.5 6 0.8) canthal scales, with 35 (4.0 6structure on truncus and lobes; no asulcate0.7) larger posterior scales; 715 (10.6 6 1.7) processus present (Fig. 4).scales present between second canthals; 1018 (13.8 6 2.1) scales present between Anolis apletophallus sp. nov.posterior canthals; 2468 (41.1 6 9.6) loreal Anolis limifrons: Bocourt (1873; in part);scales in a maximum of 58 (6.1 6 0.8)Dunn (1930; in part); Barbour (1934; inhorizontal rows, with the scales of lower rowspart); Breder (1946).and those adjacent to the canthals mostly Anolis limifrons limifrons: Etheridge (1959; inkeeled, and those of upper rows non-adjacentpart). 8. March 2008] HERPETOLOGICA99FIG. 4.Hemipenis of Anolis limifrons (SMF 85246). (a)sulcate view; (b) asulcate view. Scale bar equals 1.0 mm. Holotype (Fig. 5).SMF 85307, an adultmale from Panama City, Metropolitan Na-tional Park (8u589600N, 79u329460W), 45 m,Panama Province, Panama. Collected 26January 2006 by Gunther Kohler, 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 (SVLin largest specimen 47.0 mm) of the genusAnolis (sensu Poe, 2004) that is most similar inFIG. 5.Head of holotype of Anolis apletophallusexternal morphology to a cluster of Central (SMF 85307). Scale bars equal 1.0 mm.American species that are long-legged (lon-gest toe of adpressed hindlimb reaches tomid-eye or beyond), have a single elongated slightly imbricate (smooth and non-imbricate).prenasal scale, smooth to slightly keeled Anolis yoroensis: Ventrals weakly keeled andventral scales, and slender habitus, oftenslightly imbricate (smooth and non-imbricate).delicate (i.e., Anolis dollfusianus, A. limifrons,Anolis zeus: Hemipenis unilobed (bilobed);A. ocelloscapularis, A. rodriguezii, A. yoroen- male dewlap uniformly dull white (almostsis, A. zeus). Within this cluster of species, A. uniformly orange).apletophallus is most similar to A. limifronsDescription of the holotype.Adult male asfrom which it is readily distinguished by indicated by everted hemipenes; SVLhemipenis morphology: hemipenis small and 44.0 mm; tail length 96.0 mm, tail complete;unilobed in A. limifrons, large and bilobed intail slightly compressed in cross section, tailA. apletophallus (Fig. 6). Anolis apletophallus height 1.75 mm, tail width 1.70 mm; axilla todiffers from the remaining species in thisgroin distance 17.6 mm; head length 12.0 mm,cluster by the following characteristics (con-head length/SVL ratio 0.27; snout lengthdition for A. apletophallus in parentheses):5.2 mm; head width 7.1 mm; longest toe ofAnolis dollfusianus: Hemipenis unilobed (bi-adpressed hind limb reaching to a pointlobed); ventrals weakly keeled and slightly between eyes and rostrils; shank lengthimbricate (smooth and non-imbricate). Anolis13.0 mm, shank length/head length ratio 1.08;ocelloscapularis: Ventrals weakly keeled andlongest finger of extended forelimb reaching toslightly imbricate (smooth and non-imbricate);a point slightly beyond nostrils; longest fingeran ocellated shoulder spot present (absent).of adpressed forelimb just reaches anteriorAnolis rodriguezii: Ventrals weakly keeled andinsertion of hind limbs. Most scales on snout 9. 100HERPETOLOGICA[Vol. 64, No. 1 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 level below center of eye; 4 (right)3 (left) subocu- lars broadly in contact with supralabials; ear opening 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); 7 infralabials to level below center of eye; sublabials undifferentiated; keeled granular scales present on chin and throat; dewlap extending from level below oral ricti to 3 mm beyond level of axilla; dorsum of body with weakly keeled granular scales with rounded posterior margins, 2 medial rows slightly enlarged, largest dorsal scales about 0.21 3 0.23 mm (length 3 width); about 64 medial dorsal scales in one head length; about 85 medial dorsal scales between axilla and groin; lateral scales homogeneous, average size 0.16 mm in diameter; ventrals at midbody smooth, slightly bulging, non-imbricate, about 0.34 3 0.34 mm (length 3 width); about 42FIG. 6.Hemipenis of Anolis apletophallus (SMF ventral scales in one head length; about 6680719). (a) sulcate view; (b) asulcate view. Scale barequals 1.0 mm. ventral scales between axilla and groin; 142 scales around midbody; caudal scales stronglykeeled; 7 postrostrals; 10 scales between nasals;keeled; caudal middorsal scales slightly en-1 large elongated prenasal scale in contact with larged, without whorls of enlarged scales,both rostral and first supralabial; scales inalthough an indistinct division in segments isdistinct prefrontal depression slightly tubercu- discernible; a pair of enlarged postanal scaleslate posteriorly, wrinkled anteriorly; supraor-present, about 0.69 mm wide; no tube-likebital semicircles well developed, separated by 3 axillary pocket present; scales on dorsal surfacescales; supraorbital disc composed of 1315of forelimb keeled, imbricate, about 0.28 3distinctly enlarged keeled scales; circumorbital 0.25 mm (length 3 width); digital pads dilated;row incomplete, therefore, one enlarged su-distal phalanx narrower than and raised frompraorbital in contact with supraorbital semicir- dilated pad; 24 lamellae under phalanges iiivcles; a single large elongated superciliary; about of fourth toe; 9 (right)8 (left) scales under3 or 4 rows of small keeled scales extending distal phalanx of fourth toe.between enlarged supraorbitals and super- Description of completely everted hemipe-ciliaries; a very shallow parietal depressionnis.Medium-sized bilobed organ; sulcuspresent; interparietal scale well developed, 1.4 spermaticus bordered by well developed sulcal3 1.0 mm (length 3 width), surrounded by lips and bifurcating at base of apex; shortly afterscales of moderate size; 3 scales presentthe bifurcation, branches open into broadbetween interparietal and supraorbital semi- concave areas, one on each lobe; asulcatecircles; canthal ridge distinct, composed of 4 surface of apex and distal truncus stronglylarge (posterior two largest) and 4 smallcalyculate, base of truncus with transverseanterior canthal scales; 10 scales present folds; no asulcate processus present, althoughbetween second canthals; 17 scales present a slightly elevated ridge present (Fig. 6). 10. March 2008] HERPETOLOGICA101 Coloration in life.Dorsal ground colorWarm sepia (color 221A); flanks Dark Drab(119B); dorsal surface of head Dark Drab(119B); venter dull white suffused with DarkDrab (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 ofthe male paratypes, one pair of slightlyenlarged 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 specimenswere 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. Anolis cryptolimifrons sp. nov. FIG. 7.Head of holotype of Anolis cryptolimifrons (SMF 85230). Scale bars equal 1.0 mm.Anolis limifrons: Poe (2004; in part).Anolis limifrons limifrons: Etheridge (1959; inpart.).ventral scales, and slender habitus, often delicate (i.e., A. apletophallus, A. dollfusianus,Holotype (Fig. 7).SMF 85230, an adult A. limifrons, A. ocelloscapularis, A. rodrigue-male from Cerro Brujo (9u11916.40N,zii, A. yoroensis, A. zeus). Within this cluster82u11925.40W), 10 m, Bocas del Toro Prov-of species, A. cryptolimifrons is most similar toince, Panama. Collected 19 January 2006 by A. limifrons and A. apletophallus. Anolis Gunther Kohler, Javier Sunyer, Abel A. cryptolimifrons is readily distinguished fromBatista R. and Marcos Ponce. Field tag A. limifrons by hemipenis morphology: hemi-number GK 1502.penis small and unilobed in A. limifrons, largeParatypes.SMF 8523144, same collect- and bilobed in A. cryptolimifrons. Anolising data as holotype. Most of the paratypes arecryptolimifrons differs from A. apletophallusfemales, except SMF 8523637, 8523940,in male dewlap size and color in life (small and8524243 (adult males).dull white with a small basal orange blotch inDiagnosis.A medium-sized species (SVL N. cryptolimifrons versus dewlap large andin largest specimen 45.0 mm) of the genusalmost uniformly orange in N. apletophallus).Anolis (sensu Poe, 2004) that is most similar in Anolis cryptolimifrons differs from the re-external morphology to a cluster of Centralmaining species in this cluster by the followingAmerican species that are long-legged (lon-characteristics (condition for A. cryptolimi-gest toe of adpressed hindlimb reaches tofrons in parentheses): Anolis dollfusianus:mid-eye or beyond), have a single elongatedHemipenis unilobed (bilobed); ventrals weak-prenasal scale, smooth to slightly keeledly keeled (smooth); fewer than 80 dorsals 11. 102HERPETOLOGICA [Vol. 64, No. 1between levels of axilla and groin (more than of 4 large (posterior two largest) and 4 small87); male dewlap almost uniformly orange- anterior canthal scales; 16 scales presentyellow (dull white with a small basal orangebetween second canthals; 19 scales presentblotch). Anolis ocelloscapularis: An ocellatedbetween posterior canthals; 61 loreal scales inshoulder spot present (absent); ventrals weak-a maximum of 6 horizontal rows, with thely keeled (smooth); male dewlap almostscales of lower rows mostly keeled, and thoseuniformly orange (dull white with a small of upper rows mostly tuberculated; 6 (right) basal orange blotch). Anolis rodriguezii: Male5 (left) keeled subocular scales arranged in adewlap almost uniformly orange (dull whitesingle row; 7 supralabials to level below centerwith a small basal orange blotch). Anolis of eye; 2 suboculars broadly in contact withyoroensis: Ventrals weakly keeled (smooth); supralabials; ear opening 0.5 3 1.3 mmfewer than 87 dorsals between levels of axilla(length 3 height); mental distinctly widerand groin (more than 87); male dewlap almostthan long, completely divided medially, bor-uniformly orange (dull white with a small dered posteriorly by 8 keeled postmentalsbasal orange blotch). Anolis zeus: Hemipenis(outer pair larger); 7 infralabials to level belowunilobed (bilobed); male dewlap uniformly center of eye; sublabials undifferentiated;dull white without a basal orange blotch (basal keeled granular scales present on chin andorange blotch present). throat; dewlap extending from level below oral Description of the holotype.Adult male as ricti to about 1 mm anterior to axilla; dorsumindicated by everted hemipenes; SVL of body with weakly keeled granular scales41.0 mm; tail length 88.0 mm, tail complete;with rounded posterior margins, 2 medialtail slightly compressed in cross section, tail rows slightly enlarged, largest dorsal scalesheight 1.50 mm, tail width 1.25 mm; axilla to about 0.19 3 0.20 mm (length 3 width);groin distance 18.0 mm; head length about 54 medial dorsal scales in one head11.0 mm, head length/SVL ratio 0.27; snoutlength; about 97 medial dorsal scales betweenlength 5.2 mm; head width 6.6 mm; longest axilla and groin; lateral scales homogeneous,toe of adpressed hind limb reaching toaverage size 0.10 mm in diameter; ventrals atanterior portion of eye; shank length midbody smooth, slightly bulging, non-imbri-11.1 mm, shank length/head length ratio cate, about 0.24 3 0.27 mm (length 3 width);1.01; longest finger of extended forelimb about 46 ventral scales in one head length;reaching nostrils; longest finger of adpressedabout 68 ventral scales between axilla andforelimb not reaching anterior insertion of groin; 148 scales around midbody; caudalhind limbs. Most scales on snout keeled, some scales strongly keeled; caudal middorsal scaleswrinkled; 8 postrostrals; 10 scales between slightly enlarged, without whorls of enlargednasals; 1 large elongated prenasal scale in scales, although an indistinct division incontact with both rostral and first supralabial;segments is discernible; a pair of enlargedscales in distinct prefrontal depression slightly postanal scales present; no tube-like axillarytuberculate; supraorbital semicircles well de-pocket present; scales on dorsal surface ofveloped, separated by 2 scales; supraorbitalforelimb keeled, imbricate, about 0.24 3disc composed of 910 distinctly enlarged 0.26 mm (length 3 width); digital padskeeled scales; circumorbital row complete,dilated; distal phalanx narrower than andtherefore, no enlarged supraorbitals in contact raised from dilated pad; 24 lamellae underwith supraorbital semicircles; a single large phalanges iiiv of fourth toe on right footelongated superciliary; about 5 rows of small (fourth toe of left foot missing); 7 scales underkeeled scales extending between enlargeddistal phalanx of fourth toe.supraorbitals and superciliaries; a shallowDescription of completely everted hemipe-parietal depression present; interparietal scalenis.Medium-sized bilobed organ; sulcuswell developed, 1.30 3 0.95 mm (length 3spermaticus bordered by well developed sulcalwidth), surrounded by scales of moderate size lips and bifurcating at base of apex; shortly afteranteriorly and small size posteriorly; 3 scales the bifurcation, branches open into broadpresent between interparietal and supraorbitalconcave areas, one on each lobe; asulcatesemicircles; canthal ridge distinct, composed surface of apex and distal truncus strongly 12. March 2008] HERPETOLOGICA103relationship between the new species and itscongener Anolis limifrons Cope. Natural history notes.All type specimenswere collected active during the day in a patchof 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 blotch2 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 2a. Hemipenis unilobed Anolis limifrons _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 2b. Hemipenis bilobed Anolis cryptolimifrons _ _ _ _ _ _ _ _ _ DISCUSSION Until recently, hemipenial morphology inthe diverse group of anoles has been mostlyignored. The minority of male anoles housedin museum collections worldwide have evert-ed hemipenes. We urge future collectors ofthese anoles to attempt to fully evert thehemipenes of their specimens. Study of these FIG. 8.Hemipenis of holotype of Anolis cryptolimi-organs may reveal more cryptic species offrons (SMF 85230). (a) sulcate view; (b) asulcate view. anoles than originally envisaged by mostScale bar equals 1.0 mm.workers. This distinction is of great conserva-tional importance: whereas a large amount ofecological data is available for Anolis limifronscalyculate, base of truncus with transverse and A. apletophallus, very little is known fromfolds; no asulcate processus, although a slightly A. cryptolimifrons, a species with a relativelyelevated ridge present (Fig. 8).restricted distribution. Although A. limifrons Coloration in life.Dorsal ground colorand A. cryptolimifrons both occur in the sameRaw Umber (color 123 in Smithe 19751981) general area of mainland western Panamasuffused with Vandyke Brown (221) at mid- (Caribbean versant), no actual instance ofdorsum; venter Pale Horn Color (92); iris syntopy has been documented. Both taxaCinnamon (123A); dewlap dull white with a appear to use the same habitat and samesmall Chamois (123D) basal blotch.perching sites and might exclude each other. Variation.The paratypes agrees well with Savage (2002) reported populations ofthe holotype in general appearance, morpho- Anolis limifrons-like lizards from the slopesmetrics and scalation (see Table 1). In most ofof the volcanoes Irazu and Turrialba withthe male paratypes, one pair of slightlykeeled ventral scales but otherwise identical inenlarged postanal scales is present. Thescutellation and coloration. Boulenger (1885)female paratypes have no dewlap and nobased the description of A. godmani on thisenlarged postanal scales. characteristic, and in earlier publications Etymology.The species name cryptolimi-Savage (e.g., Savage, 1973; Savage and Villa,frons is used as a noun in apposition and 1986) recognized this form as a valid speciesreflects the similarity and suspected close and used the name A. godmani for it. More 13. 104 HERPETOLOGICA [Vol. 64, No. 1species) which has distinctly keeled ventralscales. Also, the scales on head and dorsumare strongly keeled, much more than in anyother 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 Irazuand Turrialba in Costa Rica.Acknowledgments.Collecting and exportation permitswere provided by J. Guevara Sequeira, SINAC Central,Ministerio del Ambiente y Energa (MINAE), San Jose, Costa Rica; A. Barahona, T. Garca, A. P. Martinez, E. Munoz Galeano, and C. Romero, Corporacion Hondur-ena de Desarollo Forestal (COHDEFOR), Tegucigalpa,Honduras; M. Fonseca Cuevas, S. Tijerino, B. Quintero, I.Ortega, M. G. Camacho, C. Meja, and C. Peres-Roman, 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-Universitat, Frankfurt, Germany, B. E. Sanjur, Facultadde Ciencias Naturales y Exactas de la Universidad Autonoma de Chiriqu (UNACHI), David, Panama, andJ. Vegas, Panama City, provided logistical support for ourstudies in Panama. This paper is based in part upon worksupported by the Deutscher Akademischer Austausch-dienst (DAAD) to J. Sunyer and to G. Kohler through thePartnership Program between the J. W. Goethe-Universi- tat Frankfurt am Main, Germany, und der Facultad deCiencias Naturales y Exactas der Universidad Autonomade Chiriqu (UNACHI), David, Panama. We thank L. Czupalla and J. Kohler 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 ofNatural 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-FIG. 9.Specimens in life with extended dewlaps (a) elles de Belgique (IRSNB), Bruxelles; W. E. Duellman Anolis apletophallus male (Panama City, Panama, Panama; and J. E. Simmons, University of Kansas, Natural Historynot preserved); (b) A. cryptolimifrons male (Cerro Brujo, Museum (KU), Lawrence; D. Rossman, Museum ofBocas del Toro, Panama; not preserved); (c) A. limifrons, Natural Science, Louisianna State University (LSUMZ), male (Los Algarrobos, Chiriqu, Panama; not preserved). Baton Rouge; J. Hanken and J. P. Rosado, Museum ofComparative Zoology, Harvard University (MCZ), Cam- bridge; A. Batista and M. Ponce, Museo Herpetologico derecently, Savage (2002) considered it to be anChiriqu (MHCH), David; F. Tiedemann, Naturhistor-individual variant and therefore a synonym ofisches Museum (NMW), Wien; F. Bolanos, Museo deA. limifrons. We have examined a single Zoologa Universidad de Costa Rica (UCR), San Jose; K. L. Krysko and F. W. King, Florida Museum of Naturalfemale (SMF 86924) from this general area History (UF), Gainesville; C. A. Phillips and J. Petzing,collected at 1500 m elevation (about 160 mIllinois Natural History Survey, Center for Biodiversityabove the highest Costarican record for this(UIMNH), Champaign; R. A. Nussbaum and G. Schnei- 14. March 2008]HERPETOLOGICA 105der, University of Michigan Museum of ZoologyKOHLER, G. 2001. Anfibios y Reptiles de Nicaragua. (UMMZ), Ann Arbor; R. W. McDiarmid and W. R.Herpeton, Offenbach, Germany.Heyer, National Museum of Natural History (USNM),KOHLER, G. 2003. Reptiles of Central America. Herpeton Washington, D.C.; J. Campbell and C. Franklin, TheVerlag, Offenbach, Germany.University of Texas at Arlington (UTA), Arlington; W.KOHLER, G., AND J. KREUTZ. 1999. Norops macrophallus Bohme, Zoologisches Forschungsinstitut und Museum A.(Werner, 1917), a valid species of anole from Guate-Koenig (ZFMK), Bonn; and R. Gunther, Museum fur mala and El Salvador (Squamata: Sauria: Iguanidae).Naturkunde der HumboldtUniversitat zu Berlin (ZMB),Herpetozoa 12:5765.Berlin. We thank M. Harvey and two anonymous reviewers KOHLER, G., AND J. R. MCCRANIE. 2001. Two new species for comments on an earlier version of the manuscript. of anoles from northern Honduras (Squamata: Poly-chrotidae). Senckenbergiana Biologica 81:235245. LITERATURE CITEDKOHLER, G., J. R. MCCRANIE, K. E. NICHOLSON, AND J. KREUTZ. 2003. Geographic variation in hemipenialBARBOUR, T. 1934. The anoles II. The mainland species morpholgy in Norops humilis (PETERS 1863), and the from Mexico southward. Bulletin of the Museum of systematic status of Norops quaggulus (COPE 1885) Comparative Zoology 77:119155.(Reptilia, Squamata: Polychrotidae). Senckenbergiana BOCOURT, M. F. 1873. In A. Dumerl, M. F. Bocourt, andBiologica 82:213222. F. Mocquard (Eds.), Etudes sur les reptiles. In:KOHLER, G., J. R. MCCRANIE, AND L. D. WILSON. 2001. A Recherches zoologiques pour sevir a lhistoire de lanew species of anole from western Honduras (Squa- faune de lAmerique Centrale et du Mexique. Mission mata: Polychrotidae). Herpetologica 57:247255. Scientifique au Mexique et dans lAmerique Centrale, LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. recherches Zool., part 3, sect. 1. Imprimerie au DAWSON. 1985. Standards in herpetology and ichthyol- Nationale, Paris, France.ogy: Part I. Standard symbolic codes for institutionalBOULENGER, G. A. 1885. Catalogue of the Lizards in theresource collections in herpetology and ichthyology. British Museum (Natural History). 2nd Ed., Vol. II.Copeia 1985:802832. Trustees of the British Museum, London, U.K. MEYER, J. R., AND L. D. WILSON. 1973. A distributionalBRATTSTROM, B. H., AND T. R. HOWELL. 1954. Notes onchecklist of the turtles, crocodilians, and lizards of some collections of reptiles and amphibians fromHonduras. Contributions in Science, Natural History Nicaragua. Herpetologica 10:114123.Museum, Los Angeles County 244:139.BREDER, C. M., JR.. 1946. Amphibians and reptiles of the MYERS, C. W., E. E. WILLIAMS, AND R. W. MCDIARMID. Rio Chucunaque drainage, Darien, Panama, with notes1993. A new anoline lizard (Phenacosaurus) from the on their life histories and habits. Bulletin of thehighland of Cerro de la Neblina, southern Venezuela. American Museum of Natural History 86:375436.American Museum Novitates 3070:115.COPE, E. D. 1862. Contributions to Neotropical saurology. Proceedings of the Academy of Natural Sciences of PETERS, J. A., AND R. DONOSO-BARROS. 1970. Catalogue of Philadelphia 14:176188. the Neotropical Squamata. Part II. Lizards andCOPE, E. D. 1871. Ninth contribution to the herpetology Amphisbaenians. United States National Museum of tropical America. Proceedings of the Academy of Bulletin 297:1293. PETERS, W. 1873. Uber neue Saurier (Spaeriodactylus, Natural Sciences of Philadelphia 23:200224.COPE, E. D. 1874. Description of some new species ofAnolis, Phrynosoma, Tropidolepisma, Lygosoma, reptiles obtained by Dr. John F. Bransford, assistantOphioscincus) aus Centralamerica, Mexico und Aus- surgeon, U.S. Navy, while attached to the Nicaraguan tralien. Monatsbericht der Koniglich Preussis- surveying expedition in 1873. Proceedings of the chen Akademie der Wissenschaften zu Berlin 1873: Academy of Natural Sciences of Philadelphia 26:6472.738747.COPE, E. D. 1876. On the Batrachia and Reptilia of Costa POE, S. 2004. Phylogeny of anoles. Herpetological Rica. Journal of the Academy of Natural Sciences ofMonographs 18:3789. Philadelphia 8:93154. (1875) SAVAGE, J. M. 1970. On the trail of the golden frog: WithDUNN, E. R. 1930. Notes on Central American Anolis. Warzewicz and Gabb in Central America. Proceedings Proceedings of the New England Zoological Club of the California Academy of Sciences 18:273288. 12:1524. SAVAGE, J. M. 1973. Herpetological collections made byETHERIDGE, R. 1959. The relationships of the anoles Dr. John F. Bransford, Assistant Surgeon, U.S.N. (Reptilia: Sauria: Iguanidae). An interpretation based during the Nicaragua and Panama Canal Surveys on skeletal morphology. Ph.D. Dissertation, University (18721885). Journal of Herpetology 7:3538. of Michigan, Ann Arbor, Michigan, U.S.A.SAVAGE, J. M. 1997. On terminology for the description ofFITCH, H. S., AND R. A. SEIGEL. 1984. Ecological andthe hemipenes of squamate reptiles. Herpetological taxonomic notes on Nicaraguan anoles. Milwaukee Public Journal 7:2325. Museum Contributions in Biology and Geology 57:113.SAVAGE, J. M. 2002. The amphibians and reptiles of CostaGAIGE, H. T., N. HARTWEG, AND L. C. STUART. 1937. Notes Rica. A herpetofauna between two continents, between on a collection of amphibians and reptiles from easterntwo seas. University of Chicago Press, Chicago, Illinois, Nicaragua. Occasional Papers Museum of Zoology U.S.A. University of Michigan 357:118.SAVAGE, J. M., AND J. VILLA. 1986. Introduction to theKOHLER, G. 1999. The amphibians and reptiles of Herpetofauna of Costa Rica. Society for the Study of Nicaraguaa distributional checklist with keys. CourierAmphibians and Reptiles, Contributions to Herpetolo- Forschungsinstut Senckenberg 3:1207. 15. 106HERPETOLOGICA[Vol. 64, No. 1SMITHE, F. B. 19751981. Naturalists color guide. Part I. 50535253, 50535456all H, 50535760, 505361H, 505362Color guide. 182 color swatches. American Museum of63, 505364H, 505365, 505366H, 505367, 50536870all H; Ro Natural History, New York, New York, U.S.A.Agua Salud, Panama Canal: USNM 50543234, 505435both HH HSTUART, L. C. 1955. A brief review of the Guatemalan 36, 505437, 505438 , 50543940, 505441 , 505442,lizards of the genus Anolis. Miscellaneous Publications50544345all H, 50544649, 505450H, 505451, 505452H,Museum of Zoology University of Michigan 91:131.505453, 50545455both H, 505456; Ro Gigantito, PanamaTAYLOR, E. H. 1956. A review of the lizards of Costa Rica. Canal: USNM 50551520, 50552122both H, 505523, HUniversity of Kansas Science Bulletin 38:1322. 505524 , 50552540; Ro Mendoza, Panama Canal: USNM `THOMINOT, A. 1882. Note sur un Anolis despece nouvelle. 50562842, 505643H, 50564448, 505649H; Tigre Island, Bulletin de la Societe philomatique de Paris 6:251253.Panama Canal: USNM 505484500, 50550105all H,VENCES, M., M. FRANZEN, A. FLASCHENDRAGER, R. 50550610, 505511H, 50551214; Quipo, on Ro Ciri, W SCHMITT, AND J. REGOS. 1998. Beobachtungen zurn: side L. Gatun: AMNH 42919; Darie 0.5 hr below junctionHerpetofauna von Nicaragua: Kommentierte Artenliste of Ro Jaque and Ro Imamado: USNM 161216; Pirrider Reptilien. Salamandra 34:1742. Range, near head of Ro Limon: USNM 50151; Chalichi-VILLA, J., L. D. WILSON, AND J. D. JOHNSON. 1988. Middle mans Creek: AMNH 4291617; Camp Creek: AMNHAmerican Herpetology. University of Missouri Press, 42920; Ro Chucunague [Chucunaque]: AMNH 3790203,Columbia, Missouri, U.S.A. 49214; Ro Chucunague [Chucunaque], 3 mi W of CampWILSON, L. D., AND J. R. MCCRANIE. 1994. Comments on Townsend: AMNH 10255758; Yavisa, backyard junkpiles:the occurence of a salamander and three lizard species CM 7403031; Yavisa, trail along Ro Chucunague [Chucu-in Honduras. AmphibiaReptilia 15:416421. naque]: CM 74038; 711 km SW El Real between Ro Presencia and Ro Morgentese, 100350 m: UMMZ 15580203; Ro Sucubti: AMNH 4290914; Tapia: AMNH .Accepted: 9 January 2008 25021, 25023, 2502526; Panama: Metropolitan National .Associate Editor: Michael Harvey Park, Panama City (8u589600N, 79u329460W), 45 m: SMF 85307*H, 85308*, 8530910both*H, 8531119all*; Panama Prov.: AMNH 71727, 71729, 7600103, 89883; Canal APPENDIXZone: AMNH 67078, 6708182, 71716, 7173033, 75990H, Specimens Examined.*Specimens used in Discrimi-7599192, 8988082, 8560507, 10743264, 107465H,nant Function Analysis 107466, USNM 54325; Las Cascadas, Canal Zone: MCZ H Specimens with everted hemipenis19414*, 19416*, 175185*, 175187*, 17518990both*, Anolis apletophallus.Panama: Colon: Cano Saddle, 175194*, 17519697both*, 175200*, 175203*, 175207*,Closes plantation: USNM 69586; Chagres River: USNM175209*, 17521316all*, 175223*, 175225*, 175233*; Canal 102849; Ro Indio, near Gatun: USNM 54007; Gatun: Zone, Camp Mary Caretta (5 Camp Santa Margarita):USNM 5401112; Bohio Peninsula East, Panama Canal: USNM 2516263; Canal Zone, Gamboa: USNM 193351;USNM 50523137, 50523842all H, 50524351, 505252H, Canal Zone, Ro Frijoles, 3 mi N Gamboa: UF 124417;505253, 50525456all H, 505257, 505258H, 505259, Gamboa, at confluence of Panama Canal and Chagres River505260H, 50526169, 50527073all H, 505274, 505275(9u069540N, 79u419420W): USNM 29780709; Gamboa:76both H, 50527785, 505286H, 50528788, 50528990both H, SMF 84954, 8308485; Canal Zone, Ro Medio: USNM50529192, 505293301all H; Buena Vista Peninsula, 102725; Lion Hill: USNM 54172; Old Panama: USNMPanama Canal: USNM 505199230; Gigante Ridge, Pana-50129; Panama, Cabima (Pacific slope): USNM 4850001;ma Canal: USNM 50537182, 50538384both H, 505385, Panama, Cocoli: USNM 193365, 193371, 52337778;50538688all H, 50538993, 50539495both H, 50539697, Puerta Obaldia, Quebrada Repressa: USNM 150127;50539899both H, 505400, 505401H; Juan Gallegos Island Venado Beach: USNM 193359; Punta de Pena: USNMEast, Panama Canal: USNM 50557182, 50558384both H, 38712; Toro Point: USNM 53725; Trinidad River: USNM505585, 505586H, 50558789, 505590H, 50559197, 6399294, 63997; Viento Fro: USNM 48597; La Joya,50559899both H, 505600, 505601H; Juan Gallegos Island Pacific side: ANSP 25136*; Ro Tatare, Pacific side: ANSPNorth, Panama Canal: USNM 50560204, 50560508all H, 25137; Gatun: ANSP 1952022all*, 19523, 2486365all*, 50560916, 505617H, 50561822, 505623H, 505624,2486667; Canal Zone, Fort Gulick, Atlantic side: ANSP505625H, 505626, 505627H; Juan Gallegos Island South,25104*; Canal Zone, Empire: ANSP 19545; Chico: CMPanama Canal: USNM 50554149, 505550H, 50555152,6859; Barro Colorado Island: AMNH 75986, ANSP 24487505553H, 505554, 50555557all H, 505558, 505559H,92all*, 24493*H, 24494500, 24559, CM 7664, 7666, 7669,50556063, 50556468all H, 505569, 505570H; Limbo7671, 767375, 7681, 768693, 7699, 7705, 770710, 7712Camp, Panama Canal: USNM 50545773, 505474H, 19, 7725; Barro Colorado Island, Wheeler trail: UMMZ50547579, 50548081both H, 505482, 505483H; Lion Hill 63688; Lutz Creek below Donats bridge: CM 7659; Ro Island, Panama Canal: USNM 50530205, 50530608all H,Pequeni, head of Madden Lake: ANSP 21694; Juan Mina,505309, 505310H, 50531118, 50531920both H, 50532131,Madden Lake watershed: CM 74047; near Fort Clayton505332H, 50533334; Pena Blanca Peninsula, PanamaReservation: UIMNH 42184; Cerro Campana: AMNHCanal: USNM 50540206, 505407H, 50540810, 505411 75999; Cerro Campana, 800900 m: AMNH 10666; Altos15 all H, 505416, 505417 , 50541822, 505423H, 50542425,H de Maje AMNH 109623H, 10962435; Serrana de Maje: ,50542627both H, 505428, 50542930both H, 505431; Poach- proximities of Union Saldana, Ro Chiman (8u51959.00N,ers Peninsula: USNM 50565059, 50566061both H, 78u35913.60W), 470 m: MHCH 1146; Serrana de Maje Ro, 50566264, 505665H, 50566667, 50566869both H,Ambroya: MHCH 1082H, 1086, 1090; San Blas: Armila: H505670, 50567173all H, 50567476; Puma Island, Panama USNM 150099, 150100 , 150101108; Armila, QuebradaCanal: USNM 50533544, 505345H, 50534650, 505351H,Venado: USNM 15011013; Nusagandi, near field station 16. March 2008]HERPETOLOGICA107(9u20.509N, 78u59.649W), 300360 m: SMF 80717H, 80718, mouth of Ro Tortuguero, ca. 50 mi NW Limon: AMNH 80719H , 80720*H; km 14.6 on El Llano Cart road, 370 m:89174H; ca. 5 mi N Limon: AMNH 89177H; Zent AMNH 110572; km 12.8 on El Llano Cart road, 290 m:[10u01960 N, 83u16960 W, 31 m]: USNM 137767;AMNH 11057374.Approximately 17.0 km WSW Puerto Limon between Anolis cryptolimifrons.Costa Rica: Limon: SE side Ro Blanco and Ro Toro R1288288; Siquirres largeCerro Nimaso: UCR 8477*H; Panama: Bocas del Toro:stream outside of town R1288081; Motel Matama,Cerro Brujo (9u11916.40N, 82u11925.40W), 10 m: SMF 3.5 km N Limon: AMNH 13860405; 4 mi SW La85230*H, 8523135all*, 8523637both*H, 8523841all*, Fortuna: IRSNB 11684; Atalanta Farm, Estrella Valle:8524243both*H, 85244*; vicinity of Almirante: ANSPANSP 21465; Puerto Limon: ANSP 19570, 19571H, 3404750all*H, 34051*, 34052H, 3405354, USNM1957278, IRSNB 13804, ZSM 85/1998, 86/1998; La 27906271, 27913033; Ro Changuinola, near Quebrada Castilla, lower Ro Reventazon: ANSP 2371037, 24501El Guabo, 16 km airline W Almirante, 200250 m:04, 34747; Tortuguero: UF 13578384; Tortuguero, just NAMNH 119043H, 11904449; Cayo Agua, Punta Norte: of Caribbean Conservation Commission Camp: USNMUSNM 150005, 15000709; Cayo Agua, near Punta 244861; 2.4 km E Siquirres, along Ro Pacuare: CMLimon: USNM 33869092; Isla Bastimentos, Old Point: 8956667; Rte 32, 69 km E Ro Hondo, 2 km N on dirtUSNM 29788897; Isla Bastimentos: SMF 85229, road: ANSP 32372, 32374; 23 km (air) NW Bribri at Ro85245*H; Isla Colon, ca. 0.8 mi N of Bocas del Toro Carbon along road to Uatsi: ANSP 32559; RB Hitoy (town): USNM 33821416; Isla Colon, just N of Bocas delCerere: SMF 86925*H, 86926*, 8692728both*H; EstacionToro (town), along beach at fairgrounds, E side of Biologica Tierra Media, Matina: UCR 12399*H; QuebradaH isthmus: USNM 346901; Isla Colon, La Gruta: USNM Uatsi: UCR 13031* ; Sendero San Mateo, Cerro Uatsi:31376778, 313794, 338217, 338218H, 33821922, UCR 13195*H; Finca Bryan Kubicki, Guayacan: UCR 338223H, 33822426, 33822729all H, 33823032; Isla16914*H; Puntarenas: Puntarenas Prov.: AMNH 16357;Cristobal, Bocatorito camp: USNM 348191H, 34819294, 7 mi E Golfito: LSUMZ 30260; Pennsula de Osa, Golfo348195H, 348196201, 348202H, 34820305; NW side of Dulce, Puerto Jimenez, jardin at Jimenez Yacht Club:Isla Cristobal: USNM 348206H, 34820710; Isla Pastores, SMF 8151215; EB San Gerardo, Monteverde: UCRFord Point: USNM 31384748; Isla Popa, 1 km SE of13652*H; Hotel Sunset, 1 km N Santa Elena, MonteverdeDeer Island channel: USNM 29812135; Isla Popa, southregion (10u19936.90N, 84u49924.10W), 14501475 m:end of, 1 km E of Sumwood Channel: USNM 31921325, SMF 8554953all*, 85554*H; San Jose: San Jose: ANSP 34726063, 34726465both H, 34726671, 34727275all H, 7804, USNM 8090205; San Jose, grounds of Hotel Irazu: 34727677, 347278H, 347279, 347280H, 34728183; Isla UMMZ 143761; 6.0 km N San Isidro de Perez Zeledon:Popa (9u139140N, 82u089280W), 10 m: SMF 85247 UTA R12868; Talamanca: USNM 75956; Moravia de48both*H, 85249*, 85250*H; Isla Popa, NNE beachChirripo: UMMZ 128952; Near San Isidro (9u24929.10N,(09u13924.40N, 82u06936.60W), 1020 m: SMF 85399H, 83u44906.60W), 880 m: SMF 8693335; Road from85400; Laguna de Tierra Oscura, 3.7 km S of Tigre Key: General Viejo to Santa Elena (9u20906.50N,USNM 31383851, 34846770all H, 34847180; Long Bay83u39911.40W), 650 m: SMF 86929; Honduras: Colon: Point and Flat Rock Point, between, on E side of island, Quebrada Machn (15u199100N, 85u179300W), 540 m:ca. 100 yds from beach: USNM 29781617; midpoint onUSNM 53649091, 54102629; R. B. Ro Platano, El W side of Cayo Carenero: USNM 347938; N end of Cayo Ocotillal, Cabeceras de Ro Platano (15u40.39N,Roldan: USNM 348043; Isla Solarte, 10 m: SMF 85u17.19W) 370410 m: SMF 86215H, 86216, 86217H;85251*H, 8525253both*; USNM 338552, 33855355all H, Gracias a Dios: Mocoron R4617172; confluence of Ro33855657, 338558H, 33855961, 33856263both H, Wampu and Quebrada Waskista (15u009N, 84u599W),338564, 338565H, 33856668.85 m: USNM 33018384; confluence of Ro Wampu and Anolis limifrons.Costa Rica: Costa Rica: USNM Ro Patuca (14u589N, 84u599W), 60 m: USNM 33018138334, 7040610, 81198; Parismina nivel del mar: USNMboth 82; Quebrada Waskista, 85 m: SMF 8070809 *; R. B.7544446; Colombiana: USNM 6734748; Alajuela: Ro Ro Platano, Raudal Kiplatara, (15u59.99N, 84u94.89W),Fro: USNM 19514; Pizote: UCR 998889both*H,50255 m: SMF 86172H, 8617374, 86175H, 8617682,10646*H; Pilon, Bijagua: UCR 10504*H; Laguna Lagarto8618384both H, 86186; R. B. Ro Platano, Ro CuyamelLodge, Boca Tapada: UCR 12609*H; Ro Tapezco: UCR(15u58.29N, 84u99.39W), 115345 m: SMF 86188,16515*H; Cartago: 10.0 km NE Turrialba on E bank of86189H, 86190, 86191H, 8619296, 86198H, 8620405;Ro Reventazon R1287375; 2.0 km W Pavones de R. B. Ro Platano, Pomokir (15u49.39N, 84u94.89W), 150Turrialba R1287879; 3.0 km NE Pavones de Turrialba240 m: SMF 86207, 8620809both H, 86211, 8621314both H;at Ro Chitara R1286972, R1287677; Tapanti: ZFMK R. B. Ro Platano, Crique Unawas (15u12.79N, 84u92.39W),48716; Turrialba Prov.: AMNH 6970710; Turrialba:180305 m: SMF 8622021, 862223both H, 8622428,SMF 77206, USNM 133180, 192586, 52337576; Esta- 86229H; Quebrada Waskista-Ro Wampu confluence cion Biologica Copal, Tausito, Pejibaye: UCR 16127*; (15u009N, 84u599W), 85 m: SMF 86887; Cabeceras del1 Km E La Pastora (9u58906.10N, 83u44918.90W), 1500 m: Ro Rus Rus, 190 m: SMF 86888; Olancho: confluence ofSMF 86924; Heredia: Rara Avis, Catarata (10u16.929N, Ro Wampu and Quebrada Siksatara (15u039N, 85u029W),84u02.749W), 700 m: SMF 81814, 81815H; Puerto Viejo: 95 m: USNM 330180; confluence of Ro Aner and Ro ZFMK 4872337, USNM 245041; La Selva Biological Wampu (15u049N, 85u069W), 110 m: SMF 80704, USNM Station, 2.6 km SE of Puerto Viejo de Sarapiqu: USNM 33017677; confluence of Ro Wampu and Ro Sausa50567790, 505691H, 50569295, 505696H, 505697,(15u049N, 85u069W), 100 m: SMF 80705, 8070607both*,505698H, 505699701; Finca Santiago (near La Selva): USNM 33017879; confluence of Ro Yanguay and Ro SMF 78433*; Limon: Limon Prov.: AMNH 8917173, Wampu (15u039N, 85u089W), 110 m: USNM 330175;8917576, 95095, 9967176, 14961120; ca 5/4 mi SParque Nacional Patuca, Matamoros (14u409210N, 17. 108 HERPETOLOGICA [Vol. 64, No. 185u239110W), 150 m: SMF 80710, 80712*; Parque Nacional fera Bosawas, Ayapal (13u46.619N, 85u24.149W), 195 m: Patuca, Quebrada El Guasimo (14u349380N, 85u179540W),SMF 78404*;CordilleraIsabelia (13u19.999N,140 m: SMF 80713*, 8071415, 80716H; Parque Nacional 85u41.529W), 500 m: SMF 78314H, 78315, 78316H; FincaPatuca, Caobitas (14u399220N, 85u179430W), 100 m: SMF Santa Enriqueta, 5 km E San Jose De Las Latas, 1300 m:80711; Quebrada Siksatara-Ro Wampu confluenceSMF 84732*H, JS 061*; Rait, along Ro Coco: JS 919921; (15u049N, 85u029W), 95 m: SMF 8688990; Ro Yanguay- Matagalpa: Selva Negra (12u59.969N, 85u54.559W): SMF Ro Wampu confluence (15u039N, 85u089W), 110 m: SMF7720203both*, 7720405, 77334, 77551, 78201, 78202H,8689193; Ro Aner-Ro Wampu confluence (15u039N,7820304, 78205H ,78206, 78281, 78312, 79814*H; 12 km85u079W), 110 m: SMF 8689496; Ro Sausa-Ro Wampu NE Matagalpa, 1100 m: KU 19507778; Finca Tepeyac,confluence (15u049N, 85u069W), 100 m: SMF 86897900, 10.5 km N and 9 km E Matagalpa, 960 m: KU 8565155,86901H, 8690203; Yapuwas (14u589N, 85u009W), 60 m: 85675; road to Puerto Cabezas (13u17.119N, 85u42.949W),SMF 8690406; Between Ro Sausa and Ro Wampu, 1000 m: SMF 78313; Cerro Musun, FUNDENIC 100 m: SMF 86907, 86908; Quebrada de Las Maras (12u57918.80N, 85u13951.20W), 630 m: JS 737H, 740,(15u189N, 85u219W), 660 m: SMF 86909H; Matamoros H 753 ; Ro San Juan: Machuca: ANSP 7803*; at Isla de(14u409N, 85u239W), 150 m: SMF 8691014all H; Quebrada Diamante on Ro San Juan: OMNH 3593233; Ro San El Mono (14u399N, 85u209W), 100 m: SMF 86915;Juan: USNM 2498182; San Juan del Norte: USNM 19506 Quebrada El Guasimo (15u059N,86u259W), 140 m: SMF 07; Ro San Juan, Boca de San Carlos (10u47.269N,86916; Nicaragua: no specific locality: USNM 13739,84u11.709W), 20 m: SMF 79815*, 79816H; Boca de San 15212; Palvon (5 El Polvon?): USNM 120758; Atlantico Carlos (10u47925.70N, 84u11937.70W), 40 m: JS 528; Ro Norte: Eden Mine: ANSP 21138*; Great Falls, Pispis:San Juan, Bartola (10u58.379N, 84u20.359W), 30 m: SMFANSP 21124*; Parque Nacional Saslaya, Estacion BiologicaH 79817* , 79818, 8092439, 80965; Boca de Bartola Salto Labu (13u399510N, 85u00955.50W), 260 m: SMF(10u58918.30N, 84u20923.10W), 25 m: JS 454H, 455; conflu-8206869both*; Parque Nacional Saslaya, between Estacion ence of Ro San Juan and Ro Sarapiqu: SMF 83206H, Biologica Salto Labu and Campamento El Revenido, 400 83207; Chingo Petaca (10u44950.90N, 83u50926.30W), 40 m:500 m: SMF 82236; Parque Nacional Saslaya, Campamento H SMF 83208 ; Dos Bocas de Bartola, El AlmendroI (13u439110N, 85u029200W), 720 m: SMF 82070H; Parque(10u59943.90N, 84u16937.50W), 75 m: JS 402H; Dos BocasNacional Saslaya, Campamento El Carao (13u42.799N; de Ro Indio (11u02954.80N, 83u52948.40W), 25 m: JS 542H,84u58.669W), 400 m: SMF 79611; Parque Nacional Saslaya,543, 622H; Ro Fro, FUNDEVERDE (11u04937.00N, Campamento Las Pavas (13u44.59N; 85u01.59W), 780 m:84u44955.70W), 45 m: JS 327H, 328, 330; Ro Fro,SMF 79379H; Parque Nacional Saslaya, trail from Campa- FUNDEVERDE (11u03941.80N, 84u44927.00W), 50 m: JSmento Las Pavas to Campamento El Carao: SMF 82844; 355; along Ro Papaturro, Refugio de Vida Silvestre LosParque Nacional Saslaya, trail from Campamento Las Pavas Guatuzos: JS 953H, 954, 958H; Islas de Solentiname: KU(13u44.59N; 85u01.59W) to Campamento Los Monos 17404850, SMF 77210, 77660, 7832430; Islas de(13u45.19N; 85u02.29W), 810 m: SMF 79382*H; Parque Solentiname, Isla Mancarron, Hotel Mancarron: SMFNacional Saslaya, Campamento Los Monos (13u45.19N; 82197H; Islas de Solentiname, Isla Mancarron: SMF 82198,85u02.29W), 800820 m: SMF 79161, 79591*, 79607, 82199*; Islas de Solentiname, Isla Mancarron, 1 km N 79898*; Parque Nacional Saslaya, Campamento Las Ranas, Hotel Mancarron: SMF 82200*H; Panama: Bocas del920 m: SMF 82071*; Cerro El Toro (13u42930.60N, Toro: 7.5 km airline WSW Chiriqu Grande, 10 m: AMNH85u02917.70W), 830 m: SMF 83209; Rancho Alegre 123265, 123266H, 12326768, 123269H; Rambala, near(13u39947.50N, 85u01938.90W), 285 m: SMF 84737*, JS Chiriqu Grande (08u559270N, 82u119030W), 20 m: SMF052*; Finca URACCAN (13u43944.10N, 84u53914.50W),85254H, 85255; Isla Escudo de Veraguas, West Point:145 m: SMF 84738*, JS 074*; 4 km E Bonanza, 200 m: KU USNM 34748389, 347490H, 347491; Celentine101397; Bonanza, 260 m: KU 84867, 8565657, 101398 (8u479090N, 82u119170W), 610 m: SMF 85228*H; Quebrada400; Alamikamba (13u30.089N, 84u13.649W): SMF 77552 La Gloria, near Miramar (8u59908.00N, 82u13956.40W),55; Moss, 2 km S Finca Luciana, along Ro Wawa: JS 1089, 20 m: SMF 85246*H; Boca del Ro Krikamola (8u58938.60N, 109091both H, 1113H, 1130; Krin Krin, 5 km W confluence 81u55901.70W), 7 m: MHCH 539H; Chiriqu: Chiriqu:Ro Waspuk with Ro Coco: JS 1193; Atlantico Sur: Ro ZMB 7785, 7827; ca. 6 mi below El Hato, alongEscondido, 45 mi. from Bluefields: USNM 1973334; Ro Hwy, 4000 ft: FMNH 60275; Bugabita, 100200 m:Escondido, 50 mi from Bluefields: USNM 19877; Cara de NMW 20673 (13); Boquete: ZSM 63/1989/3; David,Mono, 50 m: KU 112991; El Recreo, S side Ro Mico, 25 m: UNACHI, Jardn Botanico, near River: SMF 85260*;KU 10186567, 11298890; vicinity of Providencia Los Algarrobos (8u29939.10N, 82u26901.30W), 110 m:(11u31.799N, 84u22.929W), 100 m: SMF 77556; Boaco: SMF 85223*; Los Algarrobos (8u299250N, 82u269100W),Finca Santa Helena, Masigue, 30 km NE Camoapa, 600 m: SMF 84733*; Jinotega: Finca Berln (13u32.269N,130 m: SMF 85256*; Los Algarrobos (8u299470N,85u41.509W), 1015 m: SMF 78980*H; Cerro Kilambe La, 82u269000W), 130 m: SMF 8525759all*; Meseta deCueva (13u36911.10N, 85u42952.60W), 1025 m: JS 161*; Chorcha (8u249380N, 82u139100W), 190 m: SMF 85227*; ,Cerro Kilambe El Diamante (13u36951.30N, 85u44920.20W, Meseta de Chorcha (8u249500N,82u139060W), 240270 m: 1090 m: SMF 84739*; Cerro Kilambe (13u37906.20N, SMF 8522426all*; Cocle N El Valle de Anton, Cerro:85u43919.50W, 1015 m: SMF 84740*H; Reserva Biosfera Gaital (8u37.679N, 80u6.609W), 750 m: SMF 80779;Bosawas, ca. 0.5 km SE Pueblo Wiso (13u59.609N,Veraguas: 56 mi (via road) NW Santa Fe: AMNH85u19.600W), 200 m: SMF 7853435both*H; Reserva Bios-120003H.