Herpetologica, 64(1), 2008, 92–108E 2008 by The Herpetologists’ League, Inc.      TWO NEW SPECIES OF ANOLES FORMERLY REFER...
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94                                     HERPETOLOGICA                                   [Vol. 64, No. 1examined. In Type A ...
TABLE 1.—Selected measurements, proportions and scale characters of Anolis limifrons, A. apletophallus, and A. cryptolimif...
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March 2008]                             HERPETOLOGICA                                              101   Coloration in lif...
102                                      HERPETOLOGICA                                   [Vol. 64, No. 1between levels of ...
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Anolis apletophallus und cryptolimifrons köhler&sunyer 2008
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Anolis apletophallus und cryptolimifrons köhler&sunyer 2008

  1. 1. Herpetologica, 64(1), 2008, 92–108E 2008 by The Herpetologists’ League, Inc. TWO NEW SPECIES OF ANOLES FORMERLY REFERRED TO AS ANOLIS LIMIFRONS (SQUAMATA: POLYCHROTIDAE) GUNTHER KOHLER1,3 ¨ AND JAVIER SUNYER1,2 1 Forschungsinstitut und Naturmuseum Senckenberg, Senckenberganlage 25, 60325 Frankfurt a.M., Germany 2 ´ ´ ´n, Gabinete de Ecologıa y Medio Ambiente, Departamento de Biologıa, 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 sı 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 and of A. bransfordii was ‘‘collected by Bransfordsubtropical portions of North, Central and at Machuco (5 Machuca) on the Rıo San ´ ¨South America (Kohler, 2003; Savage, 2002). ´ Juan, Departamento Rıo San Juan, Nicara-Several new species have been described in gua.’’ In 1882, Thominot described Anolisrecent years indicating that more field work rivieri 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 in four specimens from ‘‘Guatemala’’ and threeanoles revealed substantial cryptic diversity ´ specimens 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 species Boulenger … [there] may have been a mixupAnolis (Dracontura) limifrons based on two in locality data in the Godman-Salvin collec-syntypes (now ANSP 7900–01) from ‘‘Vera- tions, one of the cotypes having been listed asgua.’’ According to Savage (1970:279), ‘‘today of Guatemala whereas it probably came fromthe old Veragua comprises the Provincias of Costa Rica,’’ and he also stated that ‘‘the ´Veraguas, Chiriquı and Bocas del Toro.’’ material [including the holotype of A. god-Barbour (1934:139) described the type locality mani] 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 Rıo ´ 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 and ´men (now ZMB 7827) from ‘‘Chiriquı.’’ A year bransfordii as synonyms of limifrons. Barbourlater, Cope (1874) described the new species (1934:140) stated that ‘‘Dunn has seen theAnolis bransfordii from ‘‘Nicaragua’’ based on types of limifrons, rivieri, pulchripes and rodriguezii and declares them all the same 3 CORRESPONDENCE: e-mail, gkoehler@senckenberg.de species.’’ The names trochilus, pulchripes, 92
  2. 2. March 2008] HERPETOLOGICA 93bransfordii 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 until meter 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 used ¨al., 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 of tip of snout to the anterior border of the orbit.the variation in hemipenial and scalation We 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 midline.to eastern Panama commonly referred to as We 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 a For 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 (1975–1981). 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 Chiriquı, ´ 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 oflizard’s tail, we placed the lizard for 1–2 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 cause RESULTSdamage 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. 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 branches digital lamellae) +0.176923 (total number ofcontinue to the tip of the lobes. In Type B (n loreal 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 body tween 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 lobes 5 0.284551 (number of medial ventral scales inare not calyculate. The sulcus spermaticus one head length) 20.173192 (number ofopens at the base of the apex. While these two medial 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 geographical 20.603442 (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 dull Zone (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 (larger distributed in the western and central portionsthan 150 mm2) that is almost uniformly of the Province Bocas del Toro, Panama, andorange. All Type B males have a small dewlap adjacent southeastern Costa Rica (Fig. 2).(smaller than 100 mm2) that is dull white with Species 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 Chiriquı, Bocas del Toro andsis (DFA) based on seven pholidotic charac- Veraguas), the bilobed form (our Species B)ters (number of medial ventral scales in one is restricted to the Caribbean versant; whereashead length; number of medial dorsal scales in the 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 dewlap mentioned 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, male coloration 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. 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. limifrons A. apletophallus A. cryptolimifrons = 50 = 25 = 20 R 21 R 24 R 10Maximum SVL = 41.5 47.0 45.0 R 43.5 46.5 42.0Tail length / SVL = 1.53–2.52 (2.21 6 0.19) 1.52–2.29 (2.03 6 0.17) 1.83–2.19 (2.07 6 0.10) R 1.60–2.30 (2.05 6 0.22) 1.82–2.32 (2.06 6 0.13) 1.87–2.08 (1.98 6 0.08)Tail diameter vertical / horizontal = 1.00–1.30 (1.16 6 0.07) 1.03–1.27 (1.14 6 0.08) 1.00–1.25 (1.15 6 0.06) R 1.00–1.27 (1.14 6 0.07) 1.00–1.25 (1.11 6 0.07) 1.00–1.17 (1.21 6 0.06)Head length / SVL = 0.25–0.28 (0.27 6 0.01) 0.25–0.29 (0.27 6 0.01) 0.25–0.29 (0.27 6 0.01) R 0.24–0.28 (0.26 6 0.01) 0.25–0.29 (0.27 6 0.01) 0.26–0.28 (0.27 6 0.01)Head length / head width = 1.44–1.80 (1.67 6 0.07) 1.54–1.84 (1.67 6 0.06) 1.59–1.77 (1.67 6 0.06) R 1.57–1.77 (1.67 6 0.06) 1.56–1.70 (1.65 6 0.04) 1.62–1.75 (1.66 6 0.04)Interparietal plate / ear = 0.79–5.20 (2.03 6 0.78) 0.71–4.12 (1.71 6 0.70) 0.54–2.29 (1.47 6 0.46) R 1.09–4.75 (2.19 6 0.98) 1.01–2.45 (1.69 6 0.44) 0.50–1.68 (1.09 6 0.39)Shank length / SVL = 0.26–0.33 (0.30 6 0.02) 0.29–0.32 (0.30 6 0.01) 0.26–0.31 (0.29 6 0.02) R 0.26–0.31 (0.28 6 0.01) 0.26–0.32 (0.30 6 0.02) 0.27–0.29 (0.28 6 0.01)Axilla–groin distance / SVL = 0.36–0.47 (0.42 6 0.03) 0.34–0.46 (0.39 6 0.03) 0.39–0.46 (0.43 6 0.02) R 0.41–0.53 (0.45 6 0.02) 0.35–0.48 (0.41 6 0.03) 0.40–0.46 (0.43 6 0.02) HERPETOLOGICASubdigital lamellae of 4th toe 20–27 (23.33 6 1.61) 21–26 (23.59 6 1.12) 23–29 (25.03 6 1.47)Number of scales between supraorbital semicircles 0–4 (1.78 6 0.84) 1–3 (2.04 6 0.50) 1–3 (2.20 6 0.61)Number of scales between interparietal plate and supraorbital semicircles 1–5 (2.68 6 0.86) 1–4 (2.72 6 0.68) 2–4 (3.27 6 0.64)Number of scales between subocular scales and supralabial scales 0 0 0Number of supralabial scales to level below center of eye 5–8 (6.48 6 0.69) 6–8 (6.76 6 0.56) 5–7 (6.73 6 0.52)Number of infralabials to level below center of eye 5–8 (6.30 6 0.70) 5–8 (6.37 6 0.57) 6–8 (6.62 6 0.56)Total number of loreals 24–68 (41.13 6 9.56) 25–74 (45.00 6 8.78) 36–69 (49.07 6 7.67)Number of horizontal loreal scale rows 5–8 (6.10 6 0.79) 5–8 (6.38 6 0.73) 6–8 (6.70 6 0.70)Number of postrostrals 5–9 (6.86 6 0.82) 6–8 (7.23 6 0.59) 6–9 (7.17 6 0.53)Number of postmentals 4–8 (6.45 6 0.91) 6–9 (7.28 6 0.88) 6–8 (7.03 6 0.87)Number of scales between nasals 7–11 (8.91 6 0.86) 8–11 (9.69 6 0.66) 9–12 (9.77 6 0.77)Number of scales between 2nd canthals 7–15 (10.59 6 1.69) 9–13 (10.61 6 1.06) 9–16 (12.00 6 1.55)Number of scales between posterior canthals 10–18 (13.77 6 2.10) 10–18 (13.23 6 1.86) 12–19 (15.21 6 1.76)Number of medial dorsal scales in one head length 38–70 (54.26 6 5.76) 42–74 (60.65 6 7.10) 46–70 (58.87 6 5.35)Number of medial ventral scales in one head length 26–58 (40.62 6 6.15) 36–60 (45.78 6 5.44) 34–50 (41.31 6 4.01) 95
  5. 5. 96 HERPETOLOGICA [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 McCranie FIG. 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.—A medium-sized speciesreasons the following taxa (respective type (snout–vent length [SVL] in largest specimenlocalities in parentheses) are clearly referable 43.5 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 (Chiriquı, 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 oldest dollfusianus, A. ocelloscapularis, A. rodrigue-available name for this species, our Species A zii, 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 Thominot white with a small basal orange blotch in A.is 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 A. in the remaining species. Additionally, A.rivieri Thominot to be a nomen dubium. limifrons differs from the species in thisInterestingly, there is no available scientific cluster by the following characteristics (con-name for either of our Species B and C. We dition for A. limifrons in parentheses): Anolistherefore describe them as new species below. dollfusianus: ventrals weakly keeled (smooth). Anolis limifrons Cope, 1862 Anolis ocelloscapularis: An ocellated shoulder spot 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 1.53–2.52 (2.17 6 0.21); tail ‘‘Irazu, Costa Rica.’’ Barbour (1834), Taylor slightly compressed in cross section, ratio tail (1956). height/tail width 1.00–1.30 (1.16 6 0.07); ratioAnolis limifrons Cope, 1862:178. Syntypes axilla to groin distance/SVL 0.36–0.53 (0.43 6 ANSP 7900–01 from ‘‘Veragua.’’ Bocourt 0.03); ratio head length/SVL 0.24–0.28 (0.26
  6. 6. March 2008] HERPETOLOGICA 97 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 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 0.41– hind limbs; scales on snout varying from0.52 (0.45 6 0.02); ratio head length/head almost non-keeled to keeled; 5–9 (6.9 6 0.8)width 1.44–1.80 (1.67 6 0.06); longest toe of postrostrals; 7–11 (8.9 6 0.9) scales betweenadpressed hind limb reaching to a point nasals; 1 large elongated prenasal scale inbetween anterior to eye and tip of snout; ratio contact with both rostral and first supralabial,shank length/SVL 0.26–0.33 (0.29 6 0.02); occasionally only in contact with rostral; scalesratio shank length/head length 0.97–1.24 (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 of teriorly, some of them keeled; supraorbitalsnout; longest finger of adpressed forelimb semicircles well developed, separated by 0–4reaches in between anterior to insertion of (1.8 6 0.8) scales; supraorbital disc composedhind limbs and slightly beyond to insertion of of 5–14 distinctly enlarged keeled scales;
  7. 7. 98 HERPETOLOGICA [Vol. 64, No. 1 to the canthals mostly tuberculated; 4–7 keeled subocular scales arranged in a single row; 5–8 (6.5 6 0.7) supralabials to level below center of eye; 2–5 suboculars broadly in contact with supralabials; ear opening medi- um-sized, ratio tympanum height/interparietal scale length 0.58–1.36 (0.86 6 0.16); mental distinctly wider than long, completely divided medially, bordered posteriorly by 4–8 (6.5 6 0.9) postmentals; 5–8 (6.3 6 0.7) infralabials to level below center of eye; sublabials undifferentiated; keeled granular scales pre- sent on chin and throat; dewlap extending from level below oral ricti to axilla, in some specimens extending 1–2 mm posterior to axilla; dorsum of body with weakly keeled granular scales (at least anteriorly) with rounded posterior margins, 2 medial rows slightly enlarged, 38–70 (54.3 6 5.8) medial dorsal scales in one head length; 70–112 (91.5 6 9.0) medial dorsal scales between axilla and groin; lateral scales homogeneous, ventrals at midbody smooth, slightly bulging, non–imbri- cate, 26–58 (40.6 6 6.1) ventral scales in one head length; 49–79 (64.6 6 5.7) ventral scales between axilla and groin; 108–157 (132.1 6 10.6) scales around midbody; caudal scales FIG. 3.—Head of Anolis limifrons (SMF 86900). Scale strongly keeled; caudal middorsal scalesbars equal 1.0 mm. slightly enlarged, without whorls of enlarged scales, although an indistinct division incircumorbital row usually incomplete, there- segments is discernible; a pair of slightlyfore, 0–3 enlarged supraorbitals in contact enlarged postanal scales usually present; nowith supraorbital semicircles; a single large tube-like axillary pocket present; scales onelongated superciliary; 3–6 rows of small dorsal surface of forelimb keeled, imbricate;keeled scales extending between enlarged digital pads dilated; distal phalanx narrowersupraorbitals and superciliaries; a very shallow than and raised from dilated pad; 20–27 (23.3parietal depression present in most speci- 6 1.6) lamellae under phalanges ii–iv ofmens; interparietal scale well developed, fourth toe; 7–11 (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 scales Description of completely everted hemipe-posteriorly; 1–5 (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 3–5 (4.0 6 structure on truncus and lobes; no asulcate0.7) larger posterior scales; 7–15 (10.6 6 1.7) processus present (Fig. 4).scales present between second canthals; 10–18 (13.8 6 2.1) scales present between Anolis apletophallus sp. nov.posterior canthals; 24–68 (41.1 6 9.6) loreal Anolis limifrons: Bocourt (1873; in part);scales in a maximum of 5–8 (6.1 6 0.8) Dunn (1930; in part); Barbour (1934; inhorizontal rows, with the scales of lower rows part); Breder (1946).and those adjacent to the canthals mostly Anolis limifrons limifrons: Etheridge (1959; inkeeled, and those of upper rows non-adjacent part).
  8. 8. March 2008] HERPETOLOGICA 99 FIG. 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 26 ¨January 2006 by Gunther Kohler, JavierSunyer, Abel A. Batista R. and Marcos Ponce.Field tag number GK 1672. Paratypes.—SMF 85308–19, same collect-ing data as holotype. SMF 85308–13 are adultmales, SMF 85314–19 are adult females. Diagnosis.—A medium-sized species (SVLin largest specimen 47.0 mm) of the genusAnolis (sensu Poe, 2004) that is most similar in FIG. 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, often slightly 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. limifrons Description 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 in tail 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 this groin 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). Anolis 13.0 mm, shank length/head length ratio 1.08;ocelloscapularis: Ventrals weakly keeled and longest 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 and insertion of hind limbs. Most scales on snout
  9. 9. 100 HERPETOLOGICA [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 42 FIG. 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 in although 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 13–15 of 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 ii–ivcles; 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 depression nis.—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 present the 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 small calyculate, 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. 10. March 2008] HERPETOLOGICA 101 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 paratype’s 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; in part.). 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, large Paratypes.—SMF 85231–44, same collect- and bilobed in A. cryptolimifrons. Anolising data as holotype. Most of the paratypes are cryptolimifrons differs from A. apletophallusfemales, except SMF 85236–37, 85239–40, in male dewlap size and color in life (small and85242–43 (adult males). dull white with a small basal orange blotch in Diagnosis.—A medium-sized species (SVL N. cryptolimifrons versus dewlap large andin largest specimen 45.0 mm) of the genus almost 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 Central maining species in this cluster by the followingAmerican species that are long-legged (lon- characteristics (condition for A. cryptolimi-gest toe of adpressed hindlimb reaches to frons in parentheses): Anolis dollfusianus:mid-eye or beyond), have a single elongated Hemipenis unilobed (bilobed); ventrals weak-prenasal scale, smooth to slightly keeled ly keeled (smooth); fewer than 80 dorsals
  11. 11. 102 HERPETOLOGICA [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 orange between second canthals; 19 scales presentblotch). Anolis ocelloscapularis: An ocellated between posterior canthals; 61 loreal scales inshoulder spot present (absent); ventrals weak- a maximum of 6 horizontal rows, with thely keeled (smooth); male dewlap almost scales 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: Male 5 (left) keeled subocular scales arranged in adewlap almost uniformly orange (dull white single 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 almost than 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; snout length; 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 to average 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 adpressed about 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; supraorbital forelimb keeled, imbricate, about 0.24 3disc composed of 9–10 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 ii–iv of fourth toe on right footelongated superciliary; about 5 rows of small (fourth toe of left foot missing); 7 scales underkeeled scales extending between enlarged distal phalanx of fourth toe.supraorbitals and superciliaries; a shallow Description of completely everted hemipe-parietal depression present; interparietal scale nis.—Medium-sized bilobed organ; sulcuswell developed, 1.30 3 0.95 mm (length 3 spermaticus 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 supraorbital concave areas, one on each lobe; asulcatesemicircles; canthal ridge distinct, composed surface of apex and distal truncus strongly
  12. 12. March 2008] HERPETOLOGICA 103 relationship between the new species and its congener Anolis limifrons Cope. Natural history notes.—All type specimens were collected active during the day in a patch of secondary forest. The lizards were spotted on branches and leaves of bushes and small trees, 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 2 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 2a. Hemipenis unilobed Anolis limifrons _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 2b. Hemipenis bilobed Anolis cryptolimifrons _ _ _ _ _ _ _ _ _ DISCUSSION Until recently, hemipenial morphology in the diverse group of anoles has been mostly ignored. The minority of male anoles housed in museum collections worldwide have evert- ed hemipenes. We urge future collectors of these anoles to attempt to fully evert the hemipenes 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 of ecological 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 color and A. cryptolimifrons both occur in the sameRaw Umber (color 123 in Smithe 1975–1981) 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 of ´ of the volcanoes Irazu and Turrialba withthe male paratypes, one pair of slightly keeled ventral scales but otherwise identical inenlarged postanal scales is present. The scutellation and coloration. Boulenger (1885)female paratypes have no dewlap and no based 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. 13. 104 HERPETOLOGICA [Vol. 64, No. 1 species) which has distinctly keeled ventral scales. Also, the scales on head and dorsum are strongly keeled, much more than in any other examined A. limifrons. Unfortunately, no data on hemipenis morpholgy nor on male dewlap coloration are available for represen- tatives of this population. Further collecting and study is needed in order to evaluate the status of the populations of A. limifrons-like lizards 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 Energıa (MINAE), San Jose, ´ ´ Costa Rica; A. Barahona, T. Garcıa, 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. Mejıa, and C. Peres-Roman, ´ Ministerio del Ambiente y los Recursos Naturales (MARENA), Managua, Nicaragua; Y. Hidalgo, Autoridad Nacional 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. Paiz and J. R. McCranie generously donated specimens for this study. M. Piepenbring, Botanisches Institut J. W. Goethe- ¨ Universitat, Frankfurt, Germany, B. E. Sanjur, Facultad de Ciencias Naturales y Exactas de la Universidad ´ ´ Autonoma de Chiriquı (UNACHI), David, Panama, and J. Vegas, Panama City, provided logistical support for our studies in Panama. This paper is based in part upon work supported by the Deutscher Akademischer Austausch- ¨ dienst (DAAD) to J. Sunyer and to G. Kohler through the Partnership Program between the J. W. Goethe-Universi- ¨ tat Frankfurt am Main, Germany, und der Facultad de Ciencias Naturales y Exactas der Universidad Autonoma ´ ´ de Chiriquı (UNACHI), David, Panama. We thank L. ¨ Czupalla and J. Kohler for providing some of the drawings used in this paper. For the loan of and/or access to specimens, 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- 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, Chiriquı, Panama; not preserved). Baton Rouge; J. Hanken and J. P. Rosado, Museum of Comparative Zoology, Harvard University (MCZ), Cam- ´ bridge; A. Batista and M. Ponce, Museo Herpetologico derecently, Savage (2002) considered it to be an ´ Chiriquı (MHCH), David; F. Tiedemann, Naturhistor-individual variant and therefore a synonym of ˜ isches Museum (NMW), Wien; F. Bolanos, Museo deA. limifrons. We have examined a single ´ Zoologıa 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 m Illinois Natural History Survey, Center for Biodiversityabove the highest Costarican record for this (UIMNH), Champaign; R. A. Nussbaum and G. Schnei-
  14. 14. March 2008] HERPETOLOGICA 105der, University of Michigan Museum of Zoology KOHLER, 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, The Verlag, 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 Humboldt–Universitat zu Berlin (ZMB), Herpetozoa 12:57–65.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 81:235–245. LITERATURE CITED KOHLER, 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 77:119–155. (Reptilia, Squamata: Polychrotidae). Senckenbergiana ´BOCOURT, M. F. 1873. In A. Dumerıl, M. F. Bocourt, and Biologica 82:213–222. 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 la new species of anole from western Honduras (Squa- ´ faune de lAmerique Centrale et du Mexique. Mission mata: Polychrotidae). Herpetologica 57:247–255. ´ 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 the resource collections in herpetology and ichthyology. British Museum (Natural History). 2nd Ed., Vol. II. Copeia 1985:802–832. 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 on checklist of the turtles, crocodilians, and lizards of some collections of reptiles and amphibians from Honduras. Contributions in Science, Natural History Nicaragua. Herpetologica 10:114–123. Museum, Los Angeles County 244:1–39.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 notes 1993. A new anoline lizard (Phenacosaurus) from the on their life histories and habits. Bulletin of the highland of Cerro de la Neblina, southern Venezuela. American Museum of Natural History 86:375–436. American Museum Novitates 3070:1–15.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 14:176–188. 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 297:1–293. ¨ PETERS, W. 1873. Uber neue Saurier (Spaeriodactylus, Natural Sciences of Philadelphia 23:200–224.COPE, E. D. 1874. Description of some new species of Anolis, Phrynosoma, Tropidolepisma, Lygosoma, reptiles obtained by Dr. John F. Bransford, assistant Ophioscincus) 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 26:64–72. 738–747.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 of Monographs 18:37–89. Philadelphia 8:93–154. (‘‘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 18:273–288. 12:15–24. 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 (1872–1885). Journal of Herpetology 7:35–38. 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 and the hemipenes of squamate reptiles. Herpetological taxonomic notes on Nicaraguan anoles. Milwaukee Public Journal 7:23–25. Museum Contributions in Biology and Geology 57:1–13. 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 eastern two seas. University of Chicago Press, Chicago, Illinois, Nicaragua. Occasional Papers Museum of Zoology U.S.A. University of Michigan 357:1–18. 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 Nicaragua—a distributional checklist with keys. Courier Amphibians and Reptiles, Contributions to Herpetolo- Forschungsinstut Senckenberg 213:1–212. gy 3:1–207.
  15. 15. 106 HERPETOLOGICA [Vol. 64, No. 1SMITHE, F. B. 1975–1981. Naturalist’s color guide. Part I. 505352–53, 505354–56all H, 505357–60, 505361H, 505362– Color guide. 182 color swatches. American Museum of 63, 505364H, 505365, 505366H, 505367, 505368–70all H; Rıo ´ Natural History, New York, New York, U.S.A. Agua Salud, Panama Canal: USNM 505432–34, 505435– both H H HSTUART, L. C. 1955. A brief review of the Guatemalan 36 , 505437, 505438 , 505439–40, 505441 , 505442, lizards of the genus Anolis. Miscellaneous Publications 505443–45all H, 505446–49, 505450H, 505451, 505452H, Museum of Zoology University of Michigan 91:1–31. 505453, 505454–55both H, 505456; Rıo Gigantito, Panama ´TAYLOR, E. H. 1956. A review of the lizards of Costa Rica. Canal: USNM 505515–20, 505521–22both H, 505523, H University of Kansas Science Bulletin 38:1–322. ´ 505524 , 505525–40; Rıo Mendoza, Panama Canal: USNM `THOMINOT, A. 1882. Note sur un Anolis d’espece nouvelle. 505628–42, 505643H, 505644–48, 505649H; Tigre Island, ´ ´ Bulletin de la Societe philomatique de Paris 6:251–253. Panama Canal: USNM 505484–500, 505501–05all H,VENCES, M., M. FRANZEN, A. FLASCHENDRAGER, R. ¨ ¨ 505506–10, 505511H, 505512–14; Quipo, on Rıo Ciri, W ´ SCHMITT, AND J. REGOS. 1998. Beobachtungen zur ¨ ´ ´n: side L. Gatun: AMNH 42919; Darie 0.5 hr below junction Herpetofauna von Nicaragua: Kommentierte Artenliste ´ ´ of Rıo Jaque and Rıo Imamado: USNM 161216; Pirri der Reptilien. Salamandra 34:17–42. ´ ´ Range, near head of Rıo Limon: USNM 50151; Chalichi-VILLA, J., L. D. WILSON, AND J. D. JOHNSON. 1988. Middle man’s Creek: AMNH 42916–17; Camp Creek: AMNH American Herpetology. University of Missouri Press, ´ 42920; Rıo Chucunague [Chucunaque]: AMNH 37902–03, Columbia, Missouri, U.S.A. ´ 49214; Rıo Chucunague [Chucunaque], 3 mi W of CampWILSON, L. D., AND J. R. MCCRANIE. 1994. Comments on Townsend: AMNH 102557–58; Yavisa, backyard junkpiles: the occurence of a salamander and three lizard species ´ CM 74030–31; Yavisa, trail along Rıo Chucunague [Chucu- in Honduras. Amphibia–Reptilia 15:416–421. naque]: CM 74038; 7–11 km SW El Real between Rıo ´ ´ Presencia and Rıo Morgentese, 100–350 m: UMMZ ´ 155802–03; Rıo Sucubti: AMNH 42909–14; Tapia: AMNH .Accepted: 9 January 2008 ´ 25021, 25023, 25025–26; Panama: Metropolitan National .Associate Editor: Michael Harvey Park, Panama City (8u589600N, 79u329460W), 45 m: SMF 85307*H, 85308*, 85309–10both*H, 85311–19all*; ‘‘Panama Prov.’’: AMNH 71727, 71729, 76001–03, 89883; Canal APPENDIX Zone: AMNH 67078, 67081–82, 71716, 71730–33, 75990H, Specimens Examined.—*Specimens used in Discrimi- 75991–92, 89880–82, 85605–07, 107432–64, 107465H,nant Function Analysis 107466, USNM 54325; Las Cascadas, Canal Zone: MCZ H Specimens with everted hemipenis 19414*, 19416*, 175185*, 175187*, 175189–90both*, ´ Anolis apletophallus.—Panama: Colon: Cano Saddle, 175194*, 175196–97both*, 175200*, 175203*, 175207*,Close’s plantation: USNM 69586; Chagres River: USNM 175209*, 175213–16all*, 175223*, 175225*, 175233*; Canal ´ ´102849; Rıo Indio, near Gatun: USNM 54007; Gatun: ´ Zone, Camp Mary Caretta (5 Camp Santa Margarita):USNM 54011–12; Bohio Peninsula – East, Panama Canal: USNM 25162–63; Canal Zone, Gamboa: USNM 193351;USNM 505231–37, 505238–42all H, 505243–51, 505252H, ´ Canal Zone, Rıo Frijoles, 3 mi N Gamboa: UF 124417;505253, 505254–56all H, 505257, 505258H, 505259, Gamboa, at confluence of Panama Canal and Chagres River505260H, 505261–69, 505270–73all H, 505274, 505275– (9u069540N, 79u419420W): USNM 297807–09; Gamboa:76both H, 505277–85, 505286H, 505287–88, 505289–90both H, ´ SMF 84954, 83084–85; Canal Zone, Rıo Medio: USNM505291–92, 505293–301all H; Buena Vista Peninsula, 102725; Lion Hill: USNM 54172; Old Panama: USNMPanama Canal: USNM 505199–230; Gigante Ridge, Pana- 50129; Panama, Cabima (Pacific slope): USNM 48500–01;ma Canal: USNM 505371–82, 505383–84both H, 505385, Panama, Cocoli: USNM 193365, 193371, 523377–78;505386–88all H, 505389–93, 505394–95both H, 505396–97, Puerta Obaldia, Quebrada Repressa: USNM 150127;505398–99both H, 505400, 505401H; Juan Gallegos Island – Venado Beach: USNM 193359; Punta de Pena: USNMEast, Panama Canal: USNM 505571–82, 505583–84both H, 38712; Toro Point: USNM 53725; Trinidad River: USNM505585, 505586H, 505587–89, 505590H, 505591–97, ´ 63992–94, 63997; Viento Frıo: USNM 48597; La Joya,505598–99both H, 505600, 505601H; Juan Gallegos Island – ´ Pacific side: ANSP 25136*; Rıo Tatare, Pacific side: ANSPNorth, Panama Canal: USNM 505602–04, 505605–08all H, 25137; Gatun: ANSP 19520–22all*, 19523, 24863–65all*, ´505609–16, 505617H, 505618–22, 505623H, 505624, 24866–67; Canal Zone, Fort Gulick, Atlantic side: ANSP505625H, 505626, 505627H; Juan Gallegos Island – South, 25104*; Canal Zone, Empire: ANSP 19545; Chico: CMPanama Canal: USNM 505541–49, 505550H, 505551–52, 6859; Barro Colorado Island: AMNH 75986, ANSP 24487–505553H, 505554, 505555–57all H, 505558, 505559H, 92all*, 24493*H, 24494–500, 24559, CM 7664, 7666, 7669,505560–63, 505564–68all H, 505569, 505570H; Limbo 7671, 7673–75, 7681, 7686–93, 7699, 7705, 7707–10, 7712–Camp, Panama Canal: USNM 505457–73, 505474H, 19, 7725; Barro Colorado Island, Wheeler trail: UMMZ505475–79, 505480–81both H, 505482, 505483H; Lion Hill 63688; Lutz Creek below Donats bridge: CM 7659; Rıo ´Island, Panama Canal: USNM 505302–05, 505306–08all H, Pequeni, head of Madden Lake: ANSP 21694; Juan Mina,505309, 505310H, 505311–18, 505319–20both H, 505321–31, Madden Lake watershed: CM 74047; near Fort Clayton505332H, 505333–34; Pena Blanca Peninsula, Panama ˜ Reservation: UIMNH 42184; Cerro Campana: AMNHCanal: USNM 505402–06, 505407H, 505408–10, 505411– 75999; Cerro Campana, 800–900 m: AMNH 10666; Altos15 all H , 505416, 505417 , 505418–22, 505423H, 505424–25, H de Maje AMNH 109623H, 109624–35; Serranıa de Maje ´: ´ ´,505426–27both H, 505428, 505429–30both H, 505431; Poach- ´ ˜ ´ ´ proximities of Union Saldana, Rıo Chiman (8u51959.00N,ers Peninsula: USNM 505650–59, 505660–61both H, ´ 78u35913.60W), 470 m: MHCH 1146; Serranıa de Maje Rıo ´, ´505662–64, 505665H, 505666–67, 505668–69both H, Ambroya: MHCH 1082H, 1086, 1090; San Blas: Armila: ´ H505670, 505671–73all H, 505674–76; Puma Island, Panama USNM 150099, 150100 , 150101–108; Armila, QuebradaCanal: USNM 505335–44, 505345H, 505346–50, 505351H, Venado: USNM 150110–13; Nusagandi, near field station
  16. 16. March 2008] HERPETOLOGICA 107(9u20.509N, 78u59.649W), 300–360 m: SMF 80717H, 80718, ´ mouth of Rıo Tortuguero, ca. 50 mi NW Limon: AMNH ´80719H , 80720*H; km 14.6 on El Llano – Cartı road, 370 m: ´ 89174H; ca. 5 mi N Limon: AMNH 89177H; Zent ´ ´AMNH 110572; km 12.8 on El Llano – Cartı road, 290 m: [10u01960 N, 83u16960 W, 31 m]: USNM 137767;AMNH 110573–74. Approximately 17.0 km WSW Puerto Limon between ´ ´ Anolis cryptolimifrons.—Costa Rica: Limon: SE side ´ ´ Rıo Blanco and Rıo Toro R12882–88; Siquirres largeCerro Nimaso: UCR 8477*H; Panama: Bocas del Toro: stream outside of town R12880–81; Motel Matama,Cerro Brujo (9u11916.40N, 82u11925.40W), 10 m: SMF ´ 3.5 km N Limon: AMNH 138604–05; 4 mi SW La85230*H, 85231–35all*, 85236–37both*H, 85238–41all*, Fortuna: IRSNB 11684; Atalanta Farm, Estrella Valle:85242–43both*H, 85244*; vicinity of Almirante: ANSP ANSP 21465; Puerto Limon: ANSP 19570, 19571H, ´34047–50all*H, 34051*, 34052H, 34053–54, USNM 19572–78, IRSNB 13804, ZSM 85/1998, 86/1998; La ´279062–71, 279130–33; Rıo Changuinola, near Quebrada ´ ´ Castilla, lower Rıo Reventazon: ANSP 23710–37, 24501–El Guabo, 16 km airline W Almirante, 200–250 m: 04, 34747; Tortuguero: UF 135783–84; Tortuguero, just NAMNH 119043H, 119044–49; Cayo Agua, Punta Norte: of Caribbean Conservation Commission Camp: USNMUSNM 150005, 150007–09; Cayo Agua, near Punta ´ 244861; 2.4 km E Siquirres, along Rıo Pacuare: CM ´Limon: USNM 338690–92; Isla Bastimentos, Old Point: ´ 89566–67; Rte 32, 69 km E Rıo Hondo, 2 km N on dirtUSNM 297888–97; Isla Bastimentos: SMF 85229, road: ANSP 32372, 32374; 2–3 km (air) NW Bribri at Rıo ´85245*H; Isla Colon, ca. 0.8 mi N of Bocas del Toro ´ ´ Carbon along road to Uatsi: ANSP 32559; RB Hitoy ´(town): USNM 338214–16; Isla Colon, just N of Bocas del Cerere: SMF 86925*H, 86926*, 86927–28both*H; Estacion ´Toro (town), along beach at fairgrounds, E side of Biologica Tierra Media, Matina: UCR 12399*H; Quebrada ´ H ´isthmus: USNM 346901; Isla Colon, La Gruta: USNM Uatsi: UCR 13031* ; Sendero San Mateo, Cerro Uatsi:313767–78, 313794, 338217, 338218H, 338219–22, UCR 13195*H; Finca Bryan Kubicki, Guayacan: UCR ´338223H, 338224–26, 338227–29all H, 338230–32; Isla 16914*H; Puntarenas: ‘‘Puntarenas Prov.’’: AMNH 16357;Cristobal, Bocatorito camp: USNM 348191H, 348192–94, ´ ´ 7 mi E Golfito: LSUMZ 30260; Penınsula de Osa, Golfo348195H, 348196–201, 348202H, 348203–05; NW side of ´ ´ Dulce, Puerto Jimenez, jardin at Jimenez Yacht Club:Isla Cristobal: USNM 348206H, 348207–10; Isla Pastores, ´ SMF 81512–15; EB San Gerardo, Monteverde: UCRFord Point: USNM 313847–48; Isla Popa, 1 km SE of 13652*H; Hotel Sunset, 1 km N Santa Elena, MonteverdeDeer Island channel: USNM 298121–35; Isla Popa, south region (10u19936.90N, 84u49924.10W), 1450–1475 m:end of, 1 km E of Sumwood Channel: USNM 319213–25, SMF 85549–53all*, 85554*H; San Jose: San Jose: ANSP ´ ´347260–63, 347264–65both H, 347266–71, 347272–75all H, ´ 7804, USNM 80902–05; San Jose, grounds of Hotel Irazu: ´347276–77, 347278H, 347279, 347280H, 347281–83; 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 beach Chirripo: UMMZ 128952; Near San Isidro (9u24929.10N,(09u13924.40N, 82u06936.60W), 10–20 m: SMF 85399H, 83u44906.60W), 880 m: SMF 86933–35; Road from85400; Laguna de Tierra Oscura, 3.7 km S of Tigre Key: General Viejo to Santa Elena (9u20906.50N,USNM 313838–51, 348467–70all H, 348471–80; Long Bay 83u39911.40W), 650 m: SMF 86929; Honduras: Colon: ´Point and Flat Rock Point, between, on E side of island, ´ Quebrada Machın (15u199100N, 85u179300W), 540 m:ca. 100 yds from beach: USNM 297816–17; midpoint on USNM 536490–91, 541026–29; R. B. Rıo Platano, El ´ ´W side of Cayo Carenero: USNM 347938; N end of Cayo ´ ´ Ocotillal, Cabeceras de Rıo Platano (15u40.39N,Roldan: USNM 348043; Isla Solarte, 10 m: SMF 85u17.19W) 370–410 m: SMF 86215H, 86216, 86217H;85251*H, 85252–53both*; USNM 338552, 338553–55all H, ´ Gracias a Dios: Mocoron R46171–72; confluence of Rıo ´338556–57, 338558H, 338559–61, 338562–63both H, ´ Wampu and Quebrada Waskista (15u009N, 84u599W),338564, 338565H, 338566–68. 85 m: USNM 330183–84; confluence of Rıo Wampu and ´ ´ Anolis limifrons.—Costa Rica: ‘‘Costa Rica’’: USNM ´ Rıo Patuca (14u589N, 84u599W), 60 m: USNM 330181–38334, 70406–10, 81198; Parismina nivel del mar: USNM both 82; Quebrada Waskista, 85 m: SMF 80708–09 *; R. B.75444–46; Colombiana: USNM 67347–48; Alajuela: Rıo ´ ´ ´ Rıo Platano, Raudal Kiplatara, (15u59.99N, 84u94.89W),Frıo: USNM 19514; Pizote: UCR 9988–89both*H, ´ 50–255 m: SMF 86172H, 86173–74, 86175H, 86176–82,10646*H; Pilon, Bijagua: UCR 10504*H; Laguna Lagarto ´ 86183–84both H, 86186; R. B. Rıo Platano, Rıo Cuyamel ´ ´ ´Lodge, Boca Tapada: UCR 12609*H; Rıo Tapezco: UCR ´ (15u58.29N, 84u99.39W), 115–345 m: SMF 86188,16515*H; Cartago: 10.0 km NE Turrialba on E bank of 86189H, 86190, 86191H, 86192–96, 86198H, 86204–05; ´ ´Rıo Reventazon R12873–75; 2.0 km W Pavones de ´ ´ R. B. Rıo Platano, Pomokir (15u49.39N, 84u94.89W), 150–Turrialba R12878–79; 3.0 km NE Pavones de Turrialba 240 m: SMF 86207, 86208–09both H, 86211, 86213–14both H; ´ ´at Rıo Chitarıa R12869–72, R12876–77; Tapanti: ZFMK ´ ´ R. B. Rıo Platano, Crique Unawas (15u12.79N, 84u92.39W),48716; ‘‘Turrialba Prov.’’: AMNH 69707–10; Turrialba: 180–305 m: SMF 86220–21, 86222–3both H, 86224–28,SMF 77206, USNM 133180, 192586, 523375–76; Esta- 86229H; Quebrada Waskista-Rıo 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: ´ Rıo Rus Rus, 190 m: SMF 86888; Olancho: confluence ofSMF 86924; Heredia: Rara Avis, Catarata (10u16.929N, ´ ´ Rıo Wampu and Quebrada Siksatara (15u039N, 85u029W),84u02.749W), 700 m: SMF 81814, 81815H; Puerto Viejo: ´ 95 m: USNM 330180; confluence of Rıo Aner and Rıo ´ZFMK 48723–37, USNM 245041; La Selva Biological ´ Wampu (15u049N, 85u069W), 110 m: SMF 80704, USNM ´Station, 2.6 km SE of Puerto Viejo de Sarapiquı: USNM ´ ´ 330176–77; confluence of Rıo Wampu and Rıo Sausa ´505677–90, 505691H, 505692–95, 505696H, 505697, (15u049N, 85u069W), 100 m: SMF 80705, 80706–07both*,505698H, 505699–701; Finca Santiago (near La Selva): ´ USNM 330178–79; confluence of Rıo Yanguay and Rıo ´ ´ ´SMF 78433*; Limon: ‘‘Limon Prov.’’: AMNH 89171–73, ´ Wampu (15u039N, 85u089W), 110 m: USNM 330175;89175–76, 95095, 99671–76, 149611–20; ca 5/4 mi S Parque Nacional Patuca, Matamoros (14u409210N,