koehler anolis tenorioensis

Accepted by S. Carranza: 17 Oct. 2011; published: 6 Dec. 2011

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2011 · Magnolia Press

Zootaxa 3120: 29–42 (2011) www.mapress.com/zootaxa/ Article

29

A new species of anole related to Anolis altae from Volcán Tenorio, Costa Rica (Reptilia, Squamata, Polychrotidae)

GUNTHER KÖHLERSenckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25, 60325 Frankfurt a.M., Germany. E-mail: [email protected]

Abstract

I describe a new species of Anolis related to Anolis (or Norops) altae from Volcán Tenorio, Costa Rica. The new speciesdiffers from all other species in the A. altae complex by having the ventral surfaces of body and limbs distinctly reticulatedwith dark brown pigment and a male dewlap that is dark red with brown blotches, as well as in several morphometric andpholidotic characteristics. It further differs from its geographically nearest congener of this complex, A. monteverde, byhaving a bilobate hemipenis (unilobate in A. monteverde).

Key words: Anolis, Costa Rica, new species, Polychrotidae, Reptilia, Squamata, Volcán Tenorio

Resumen

Se describe una nueva especie de Anolis relacionada con Anolis (o Norops) altae, proveniente de Volcán Tenorio, CostaRica. La nueva especie difiere de todas las demás especies del complejo A. altae en tener la superficie ventral del cuerpoy extremidades reticuladas con pigmento marrón, y los machos con papera rojo oscuro manchada de marrón, así comootras características morfométricas y de escamación. Además, se diferencia de su congénere geográficamente más cercanode este complejo, A. monteverde, por tener un hemipene bilobado (unilobado en A. monteverde).

Introduction

The highlands of Lower Central America are home to a distinct assemblage of beta anoles (fide Etheridge 1959)that mainly belongs to two radiations, species related to Anolis altae (i.e., A. altae, A. fortunensis, A. gruuo, A.kemptoni, A. monteverde, A. pseudokemptoni) and those related to A. pachypus (i.e., A. magnaphallus, A. pachypus,A. pseudopachypus, A. tropidolepis and another species in the process of being described by Lotzkat et al. in press).The species in each of these species clusters show an allopatric distribution pattern and, therefore, at any givenlocality only one representative of each cluster is found. In the Serranía de Tabasará, A. pseudopachypus occurssyntopically either with A. pseudokemptoni, A. gruuo, or A. fortunensis; in the eastern portion of the Cordillera deTalamanca in western Panama and eastern Costa Rica A. kemptoni is found in sympatry either with A. magnaphal-lus or with A. pachypus, depending on the locality; in the western portion of the Cordillera de Talamanca, as well asin the Cordillera Central of Costa Rica, A. altae occurs together with A. tropidolepis; and in the Cordillera deTilarán, A. monteverde can be found with A. tropidolepis (Köhler et al. 2007, Köhler 2008). Aside from these twoclusters, only a few other species of beta anoles are commonly found above 1500 m elevation including A. datzo-rum, A. laeviventris, A. salvini, and A. woodi.

A single female specimen from Cerro Cacao referred to the A. altae complex by Köhler (2009) is the onlyrepresentative of this species complex known from the Cordillera de Guanacaste. This record in the far northwes-tern corner of the country makes it appear as likely that A. altae-like populations occur on the other peaks of thismountain range. In March 2011, together with several local naturalists, I undertook an expedition to Volcán Teno-rio, which is the southeasternmost peak in the Cordillera de Guanacaste reaching an altitude of 1916 m above sea

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level. We climbed the northern slope of this volcano starting ca. 3 km north of the town of Bijagua (i.e., at Helico-nias Lodge at 720 m elevation). In the cloud forest at an elevation of 1160 m above sea level, we collected a maleand two females belonging to the A. altae complex. However, it was immediately evident that these anoles are notconspecific with any of the known species of anoles and, therefore, I describe it as a new species below.

Material and methods

A list of the comparative specimens examined is provided in the Appendix. Abbreviations for museum collectionsfollow those of Leviton et al. (1985). Nomenclature of scale characters follows that of Köhler (2008). Scale sizeswere measured using the ocular micrometer of a stereo microscope (Leica MZ 12) and rounded to the nearest 0.01mm. All other measurements were made using precision calipers and were rounded to the nearest 0.1 mm. Headlength was measured from the tip of the snout to the anterior margin of the ear opening. Snout length was measuredfrom the tip of the snout to the anterior border of the orbit. Head width was determined as the distance between theoral ricti. Dorsal and ventral scales were counted at midbody along the midline. Tail height and width were meas-ured at the point reached by the heel of the extended hind leg. Subdigital lamellae were counted on phalanges ii toiv of the 4th toe. I considered the scale directly anterior to the circumnasal to be a prenasal. The capitalized colorsand color codes (the latter in parentheses) are those of Smithe (1975–1981). Abbreviations used are 2Canths(number of scales between second canthals), DHL (number of medial dorsal scales in one head length), HL (headlength), HW (head width), INL (infralabials), IP (interparietal plate), SAM (scales around midbody), SO (subocu-lar scales), SPL (supralabial scales), SS (supraorbital semicircles), SVL (snout–vent length), and VHL (number ofmedial ventral scales in one head length).

Results

Anolis tenorioensis sp. nov.Figs. 1, 2a, 3, 4, 5, 6

Holotype. SMF 91985, an adult male from near Laguna Danta, Volcán Tenorio (10.70521°N, 85.03068°W,WGS84), 1160 m, Alajuela Province, Costa Rica; collected 5 March 2011 by Gunther Köhler, Donald Varela Soto,and Joséph Vargas. Field tag number GK 3324.

Paratypes. SMF 91986, UCR 21182, same collecting data as holotype. Diagnosis. A medium-sized species (SVL in largest specimen 45.0 mm) of the genus Anolis (sensu Poe 2004)

that differs from all Lower Central American anoles except Anolis altae, A. carpenteri, A. fortunensis, A. fuscoau-ratus, A. gruuo, A. kemptoni, A. monteverde and A. pseudokemptoni in that it is short-legged (longest toe ofadpressed hind leg reaches only to ear opening), has smooth to slightly keeled ventral scales, and a slender habitus,often delicate. Anolis tenorioensis differs from all the species in the cluster listed above by having the ventral sur-faces of body and limbs distinctly reticulated with dark brown pigment (versus immaculate dirty white or creamcolored); and a male dewlap that is dark red with brown blotches (versus more or less uniformly reddish orange inA. altae and A. monteverde, uniform orange in A. carpenteri, posterior portion of male dewlap greenish beige, ante-rior portion reddish orange in A. fortunensis, uniform pinkish brown to red in A. fuscoauratus, more or less uniformdull orange in A. gruuo, posterior portion rose pink, anterior portion burnt orange, small cream colored basal blotchin some populations in A. kemptoni, posterior portion vinaceous, anterior portion orange, anterodorsal corner creamcolor in A. pseudokemptoni); and by having a divided prenasal scale (versus usually a single elongated prenasalscale). It also differs from A. fortunensis, A. kemptoni, and A. monteverde by having a bilobate hemipenis (versusunilobate). Additionally, it can be differentiated from A. carpenteri by having a brownish dorsal ground color (ver-sus greenish). Anolis tenorioensis is most similar to A. altae and A. monteverde, from which it further differs in sev-eral morphometric and pholidotic characteristics, such as relative head length (ratio head length / SVL) and numberof scales around midbody (see Table 1).

Description of the holotype. Adult male as indicated by everted hemipenes and presence of large dewlap (Fig.3); SVL 42.0 mm; tail length 36.0 mm, tail incomplete; tail round in cross section, tail height and width 1.9 mm;axilla to groin distance 18.7 mm; head length 10.7 mm, head length/SVL ratio 0.255; snout length 4.9 mm; head


Zootaxa 3120 © 2011 Magnolia Press · 31ANOLIS TENORIOENSIS COSTA RICA

FIGURE 1. Anolis tenorioensis in life. (A) SMF 91985 (male holotype); (B) SMF 91986 (female paratype); (C) UCR 21182(female paratype). Photos by Gunther Köhler.



FIGURE 2. Dewlaps of males in life of (A) Anolis tenorioensis (SMF 91985); (B) A. gruuo (SMF 85416); (C) A. kemptoni;(D) A. pseudokemptoni (SMF 85420); (E) A. altae (SMF 85555); (F) A. monteverde (SMF 85541). Photos by Gunther Köhler.

width 6.1 mm; longest toe of adpressed hind limb reaching to tympanum; shank length 9.9 mm, shank length/headlength ratio 0.93; longest finger of extended forelimb reaching nostril; longest finger of adpressed forelimb reachesa point 3.6 mm in front of anterior insertion of hind limbs. Most scales on snout keeled (Fig. 4); 6 postrostrals; 7scales between nasals; 2 prenasal scales, the lower one in contact with both rostral and first supralabial; scales indistinct prefrontal depression slightly wrinkled; supraorbital semicircles well developed, separated by a minimumof 1 scale; supraorbital disc composed of 6–7 distinctly enlarged keeled scales; circumorbital row incomplete,therefore, one enlarged supraorbital in contact with supraorbital semicircles; two elongated, strongly overlappingsuperciliaries, the anterior one much larger than the posterior one; about 3 or 4 rows of small keeled scales extend-ing between enlarged supraorbitals and superciliaries; parietal depression absent; interparietal scale well devel-oped, 2.0 x 1.2 mm (length x width), surrounded by scales of moderate size; 3 scales present between interparietaland supraorbital semicircles; canthal ridge distinct, composed of 3 large (posterior two largest) and 5 small anterior



FIGURE 3. Holotype of Anolis tenorioensis (SMF 91985). SVL = 42.0 mm. Photos by Gunther Köhler.

canthal scales; 7 scales present between second canthals; 7 scales present between posterior canthals; 30 (right)–28(left) mostly keeled loreal scales in a maximum of 5 horizontal rows; 6 keeled subocular scales arranged in a singlerow; 7 supralabials to level below center of eye; 5 suboculars broadly in contact with supralabials; ear opening 0.7x 0.9 mm (length x height); mental distinctly wider than long, almost completely divided medially, bordered poste-riorly by 6 postmentals; 7 (right)–9 (left) infralabials to level below center of eye; sublabials undifferentiated;keeled granular scales present on chin and throat; dewlap extending from level below posterior margin of eye tolevel of axilla; dorsum of body with weakly keeled imbricate scales with rounded posterior margins, medial rowsnot enlarged, largest dorsal scales about 0.34 x 0.31 mm (length x width); about 47 medial dorsal scales in one headlength; about 72 medial dorsal scales between axilla and groin; lateral scales keeled, granular and homogeneous,average size 0.15 mm in diameter; ventrals at midbody smooth, slightly bulging, subimbricate, about 0.34 x 0.29mm (length x width); about 48 ventral scales in one head length; about 74 ventral scales between axilla and groin;101 scales around midbody; caudal scales keeled except ventrally at base of tail; caudal middorsal scales notenlarged, without whorls of enlarged scales, although an indistinct division in segments is discernible; a pair of



FIGURE 4. Head of holotype of Anolis tenorioensis (SMF 91985). Upper lateral view, middle dorsal view, lower ventral viewof chin. Scale bars equal 1.0 mm. Drawings by Gunther Köhler.



FIGURE 5. Hemipenis of Anolis tenorioensis (SMF 91985). Asulcate view above, sulcate view below. Scale bar equals 1.0mm. Photos by Gunther Köhler.

slightly enlarged postcloacal scales present, about 0.60 mm wide; no tube-like axillary pocket present; scales ondorsal surface of forelimb keeled, imbricate, about 0.21 x 0.31 mm (length x width); digital pads dilated, dilatedpad about 3 times width of non-dilated scales on distal phalanx; distal phalanx narrower than and raised fromdilated pad; 24 (right)–26 (left) lamellae under phalanges II–IV of fourth toe; 8 scales under distal phalanx offourth toe.

The completely everted hemipenis is a large bilobate organ (Fig. 5); sulcus spermaticus bordered by well



developed sulcal lips and bifurcating at base of apex into branches that continue to tips of lobes; a weak asulcateridge present; apex strongly calyculate, truncus with transverse folds.

FIGURE 6. Ventral coloration of female paratypes (left: UCR 21182; right: SMF 91986) of Anolis tenorioensis before preser-vation. Photo by Gunther Köhler.

The extended dewlap in life (Fig. 2a) has six horizontal gorgetal-sternal rows with a declining number of scalesper row from base to tip (17–19 scales per row on basal portion, 5–9 towards tip); about 50 scales in marginalseries.

Coloration in life was recorded as follows: Dorsal surface of head Burnt Umber (22); dorsal ground color ofbody and limbs Grayish Horn Color (91), with shades of Buff (124) and some Dark Brownish Olive (129) pigmen-tation along vertebral line and indistinct Dark Brownish Olive (129) crossbands on limbs; tail Straw Yellow (56)with Ground Cinnamon (239) bands; venter Buff (124) with Dark Brownish Olive (129) speckling; chin Buff (124)laterally and dirty white with Dark Brownish Olive (129) reticulum medially; iris Mikado Brown (121C); dewlapPratt’s Ruby (210) with Vandyke Brown (121) blotches.

Coloration after 4 months preservation in 70% ethanol was recorded as follows: Dorsal ground color grayishbrown with indistinct paler shadings, a cream colored temporal blotch, a dark brown interorbital bar, and faintcrossbands on limbs and tail; ventral surfaces of head, body, limbs, and tail dirty pale grayish cream with a distinctdark brown reticulum; dewlap reddish (except damaged basal area) with dark brown and dirty white gorgetals, andwith dark brown blotches, mostly around gorgetals.

Variation. The paratypes agree well with the holotype in general appearance, morphometrics and scalation(see Table 1). Coloration in life of an adult female paratype (GK 3326) was recorded as follows: Dorsal groundcolor Raw Umber (123) with shades of Straw Yellow (56), a Mars Brown (223A) interorbital bar; snout VeronaBrown (223B), and a few Dark Brownish Olive (129) markings along vertebral line; tail with Verona Brown(223B) crossbands; venter Cream Color (54) with a paramedian zone of Sepia (219) reticulation; iris Verona Brown(223B); dewlap Pale Horn Color (92) with a few Grayish Horn Color (91) gorgetals.



TABLE 1. Selected measurements, proportions and scale characters of Anolis altae, A. monteverde, and A. tenorioensis (speci-mens of Anolis altae and A. monteverde listed in Appendix I). Range is followed by mean value and one standard deviation inparentheses. For abbreviations see text.

Etymology. The name tenorioensis is derived from Volcán Tenorio where the type series of the species wascollected and to where it is probably restricted, and the Latin suffix -ensis (denoting place or locality).

A. altae♂ 26♀ 43

A. monteverde♂ 8♀ 8

A. tenorioensis♂ 1♀ 2

Maximum SVL ♂ 48.5 mm 45.5 mm 42.0 mm

♀ 50.2 mm 47.0 mm 45.0 mm

Tail length / SVL ♂ 1.89–2.30 (2.11±0.12) 1.81–1.93 (1.81±0.05) —

♀ 1.81–2.22 (2.02±0.16) 1.65–1.93 (1.78±0.12) 1.64–1.69

Tail diameter vertical / horizontal ♂ 1.04–1.64 (1.30±0.16) 1.05–1.43 (1.22±0.11) 1.00

♀ 0.95–1.40 (1.20±0.18) 1.11–1.31 (1.24±0.09) 1.13–1.33

HL / SVL ♂ 0.27–0.29 (0.28±0.01) 0.27–0.29 (0.28±0.01) 0.25

♀ 0.25–0.28 (0.27±0.01) 0.25–0.28 (0.27±0.01) 0.24–0.25

HL / HW ♂ 1.56–1.82 (1.67±0.07) 1.63–1.74 (1.69±0.04) 1.75

♀ 1.50–1.78 (1.65±0.11) 1.60–1.75 (1.69±0.07) 1.70–1.82

IP / ear ♂ 1.26–6.86 (2.73±1.72) 1.71–3.67 (2.61±0.60) 3.81

♀ 0.82–5.81 (2.73±1.98) 2.01–4.40 (2.95±1.05) 1.60–4.00

Shank length / SVL ♂ 0.20–0.25 (0.23±0.01) 0.22–0.24 (0.22±0.08) 0.24

♀ 0.18–0.25 (0.21±0.03) 0.20–0.22 (0.21±0.01) 0.21–0.22

Axilla–groin distance / SVL ♂ 0.34–0.46 (0.42±0.03) 0.38–0.46 (0.43±0.03) 0.45

♀ 0.40–0.52 (0.46±0.05) 0.40–0.49 (0.44±0.03) 0.38–0.47

DHL ♂ 44–74 (57.83±7.71) 49–78 (62.13±9.54) 47

♀ 40–66 (50.72±10.60) 49–68 (55.5±8.44) 47–48

VHL ♂ 34–56 (43.52±5.53) 47–61 (54.75±4.43) 48

♀ 24–48 (34.70±9.25) 40–52 (46.75±5.34) 43–46

SAM ♂ 122–150 (136.33±7.54) 140–156 (148.25±5.90) 101

♀ 130–168 (141.63±15.50) 138–160 (148.25±9.63) 108–126

Subdigital lamellae of 4th toe (Phalanges II–IV)

20–28 (23.11±1.63) 23–28 (25.31±1.54) 21–25

Number of scales between SS 0–3 (1.46+±0.70) 1–3 (1.78±0.48) 1–2

Number of scales between IP and SS 1–4 (2.13±0.87) 2–3 (2.33±0.45) 2–3

Number of scales between SO and SPL 0 0 0

Number of SPL to level below center of eye 5–8 (6.80±0.68) 7–9 (7.50±0.63) 6–7

Number of INL to level below center of eye 5–8 (6.43±0.79) 7–8 (7.63±0.50) 6–7

Total number of loreals 24–58 (41.73±8.55) 37–68 (47.81±8.41) 27–28

Number of horizontal loreal scale rows 4–8 (6.07±0.86) 6–9 (6.63±0.81) 5

Number of postrostrals 4–8 (6.27±0.78) 5–9 (7.06±1.06) 6

Number of postmentals 4–7 (5.60±0.74) 4–6 (5.63±0.81) 5–6

Number of scales between nasals 5–10 (7.72±1.03) 5–10 (8.25±1.39) 7

Number of scales between 2nd canthals 5–11 (8.63±1.51) 6–9 (7.81±0.98) 7

Number of scales between posterior canthals 7–15 (9.80±1.70) 7–11 (9.63±1.26) 7–10



FIGURE 7. View of the cloud forest at Volcán Tenorio (taken at a point about 1000 m elevation), Costa Rica. Photo by DonaldVarela Soto.



FIGURE 8. Habitat at the type locality of Anolis tenorioensis at Volcán Tenorio, Costa Rica. Photo by Donald Varela Soto.

Natural history notes. All type specimens were collected at night while the lizards were sleeping on low veg-etation (mostly on fern leaves) within the forest (Premontane Rainforest fide Savage 2002) near an open area that isflooded most of the year and known locally as Laguna Danta (see also Figs. 7 and 8). At the type locality, a recentlyfallen tree had created a hole in the canopy allowing sun to reach the ground. No daytime searches were done.

Geographic distribution and conservation. As currently known, Anolis tenorioensis is restricted to the high-lands of Volcán Tenorio in the Cordillera de Guanacaste, Costa Rica. It occurs in a protected area (Parque NacionalVolcán Tenorio). Only three specimens of this species are known to science. However, given its presumably smallgeographic range, it seems justified to classify A. tenorioensis as Critically Endangered based on criterion B.1.a. ofthe IUCN Red List Categories and Criteria (IUCN 2011).

Discussion

The anole species related to Anolis altae (i.e., A. altae, A. fortunensis, A. gruuo, A. kemptoni, A. monteverde, A.pseudokemptoni, A. tenorioensis) form a cluster of morphologically very similar species. Among these species, twospecies pairs are evident (i.e., A. altae / A. monteverde; A. kemptoni / A. pseudokemptoni), the species in these pairsbeing almost identical in external morphology including male dewlap coloration, and differing only in hemipenialmorphology (Köhler et al. 2007, Köhler 2009). Interestingly, the variation in hemipenial morphology of these fourspecies follows an alternating pattern in a southeastern-to-northwestern direction, as follows (Fig. 9): A. pseudo-kemptoni (bilobate hemipenis), A. kemptoni (unilobate hemipenis), A. altae (bilobate hemipenis), and A. monte-verde (unilobate hemipenis). With its bilobate hemipenis, this pattern is continued by the species described herein.The underlying evolutionary mechanisms that drive this peculiar interspecific pattern in hemipenial variationremain unknown. Prezygotic isolating mechanisms come to mind with the genital differences serving as a possiblemeans of preventing copulation between individuals of different species. However, these species have no docu-mented contact zones and their climatic and vegetational requirements make it unlikely that any of these species



currently occur in sympatry or even parapatry. This situation might have been different in past geologic times whena generally colder climate might have forced the species to inhabit a lower vertical range making contacts betweenthese species more likely. In case yet another undescribed species of the A. altae complex will be discovered onVolcán Miravalles, the next Volcáno to the northwest, its males can be expected to have a unilobed hemipenis. Onlycontinued field work will tell.

FIGURE 9. Map indicating known collecting sites of Anolis altae (circles), A. kemptoni (triangles), A. monteverde (squares),A. pseudokemptoni (star), and A. tenorioensis (diamond), with illustration of hemipenis of each species. Each symbol can repre-sent one or more nearby localities. Areas above 750 and 1800 m are shaded medium gray and dark gray, respectively.

Acknowledgments

Collecting and exportation permits were provided by Javier Guevara Sequeira, SINAC Central, Ministerio delAmbiente y Energía (MINAE), San José, Costa Rica. For the loan of and/or access to specimens I thank LindaFord, Christopher J. Raxworthy and Darrell R. Frost, American Museum of Natural History (AMNH), New York;Alan Resetar, Field Museum of Natural History (FMNH), Chicago; William E. Duellman and John E. Simmons,University of Kansas, Natural History Museum (KU), Lawrence; Jeff Seigel, Natural History Museum of LosAngeles County (LACM), Los Angeles; Douglas Rossman, Museum of Natural Science, Louisianna State Univer-sity (LSUMZ), Baton Rouge; James Hanken and José P. Rosado, Museum of Comparative Zoology, Harvard Uni-versity (MCZ), Cambridge; Victor H. Tejera N., Museo de Vertebrados, Universidad de Panama (MVUP), PanamaCity; Janalee P. Caldwell and Laurie J. Vitt, Oklahoma Museum of Natural History (OMNH), Norman; FedericoBolaños, Museo de Zoología, Universidad de Costa Rica (UCR), San José; Ronald A. Nussbaum and Greg Schnei-der, University of Michigan Museum of Zoology (UMMZ), Ann Arbor; and Ron W. McDiarmid and W. RonaldHeyer, National Museum of Natural History (USNM), Washington, D.C. For field assistance, I thank Henry BrenesGonzalez, Marlon Calderón Brenes, José Francisco Salas Vega, Donald Varela Soto, Jorge Luis Soto Obando, andJoséph Vargas. Sebastian Lotzkat, Frankfurt, and Larry David Wilson, Miami, read early drafts of the manuscriptand provided helpful comments.



References

Etheridge, R. (1967) Lizard caudal vertebrae. Copeia, 1967, 699–721.IUCN (2011) http://www.iucnredlist.org/technical-documents/categories-and-criteria.Köhler, G. (2008) Reptiles of Central America. 2nd edition. Herpeton Verlag, Offenbach, Germany.Köhler, G. (2009) A new species of Anolis formerly referred to as Anolis altae from Monteverde, Costa Rica (Squamata: Poly-

chrotidae). Journal of Herpetology, 43, 11–20.Köhler, G., Ponce, M., Sunyer, J. & Batista, A. (2007) Four new species of anoles (genus Anolis) from the Serranía de Tabasará,

west-central Panama (Squamata: Polychrotidae). Herpetologica, 63, 375–391.Leviton, A.E., Gibbs, R.H. jr., Heal, E., & Dawson, C.C. (1985) Standards in herpetology and ichthyology: part I. Standard

symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia, 1985, 802–832.Lotzkat, S., Bienentreu, J.-F., Hertz, A. & Köhler, G. (in press) A new species of Anolis (Squamata: Iguania: Polychrotidae) for-

merly referred to as A. pachypus from the Cordillera de Talamanca of western Panama and adjacent Costa Rica. Zootaxa.Poe, S. (2004) Phylogeny of anoles. Herpetological Monographs, 18, 37–89.Savage, J.M. (2002) The amphibians and reptiles of Costa Rica. A herpetofauna between two continents, between two seas.

University of Chicago Press, Chicago, Illinois, U.S.A., and London, U.K.Smithe, F.B. (1975–1981) Naturalist’s color guide. Part I. Color guide. 182 color swatches. American Museum of Natural His-

tory, New York, New York, U.S.A.

APPENDIX. Specimens examined.

Anolis altae—Costa Rica: Alajuela: 1 km E Zapote at bridge of Río Tapezco, 10°13’35’’N, 84°24’29’’W, 1680 m: LACM151320, SMF 86921–86923; Cinchona, 1600 m: KU 103953–103966; between La Cinchona and Angel Falls, 1370 m:LACM 151319; Río Angel, near Cariblanco: UCR 4348; Río Santiaguito, 10°13’N, 84°10’W, near Cinchona: UCR 4350;Río La Paz Grande, 2 airline km S Cinchona: UCR 4359; Cartago: Volcán Turrialba, 2000 m: KU 102409; Coliblanco (Eslope of Volcán Irazú): FMNH 2646; 1 km E Coliblanco, 9°57’16.6’’N, 83°46’18.4’’W, 1630 m: SMF 86917–86918; Vol-cán Turrialba S slope, 5000 ft [1524 m]: UMMZ 117745 (1–3); Río Birris about 1 km W of Capellades, 1590 m: LACM149054–149068; 1 km E Pacayas, 1600 m: LACM 149069; Río Playas where it crosses the Pacayas–Santa Cruz road,1650 m: LACM 149072, 149075; Volcán Turrialba, 8 km SE of Lechería Central, 2 km NW Pastora W Río Jesus Maria,2012 m: LACM 151321, 151324–151327; Lechería Palmira, 8.4 km SE of Lechería Central on road to Santa Cruz, 1.5 kmNW Pastora W Río Jesus Maria: LACM 151322–151323; Tapantí [9°46’N, 83°46’W], 1200 m: KU 103932–103944; 5.3mi NE Bridge at Tapantí, 1250 m: LACM 114176–85; Tapantí, bridge of Río Grande de Orosí: UCR 813; Tapantí, Que-brada Valverde: UCR 2612, 2614, 2617; Parque Nacional Tapantí, Casa de Guardabosques: UCR 9484–9488, 9493;Parque Nacional Tapantí, Sendero Oropendola: UCR 11107, 11190; Parque Nacional Tapantí, Río La Mina: UCR 11322;Parque Nacional Tapantí, Sendero La Pava: UCR 11544; Peña Blanca, Río Sordi, near Cachí: UCR 11506; 2.5 km ECachí: UCR 11508; Heredia: Volcán Barba, near the Acosta Farm, 2133 m: MCZ 29385; Rama Sur Río Las Vueltas, 2100m: KU 103911; Paso Llano, S slope Volcán Barba, 2230 m: KU 66933; 2.5 km N Vara Blanca, 1900 m: KU 66934; 3 kmon road N Vara Blanca, 10°10’55.4’’N, 84°9’10.2’’W, 1675 m: SMF 85555; Vara Blanca: KU 129336–129337; Zona Pro-tectora La Selva, 2050 m: LACM 149074; Cerro Dantas, N of Cerro Chompipe, 10°05’N, 84°04’W: UCR 8586, 15840,16061; San José: La Palma: KU 40634; ca. 5 km NW San José: AMNH 90401; Alto La Palma, 10°03’N, 83°59’W: LACM114187, UCR 2952, 7124, 7129; 2 km N La Palma on road La Palma–La Hondura, 1500 m: LACM 151309; 0.5 mi S AltoLa Palma, 1500 m: LACM 151310; 1.4 mi S Alto La Palma, 1500 m: LACM 151311H; Río Cascajal, 10°02’N, 83°57’W:UCR 6740; 11.7 mi [18,8 km] N of San Isidro del General, 6800 ft [2072 m]: UMMZ 117749; 18.5 km N San Isidro delGeneral, 1700 m: KU 103946, 103947–103952; Santa Eduviges, 9°29’10.2’’N, 83°45’18.3’’W, 1507 m: SMF 86919.

Anolis cf. altae—Costa Rica: Guanacaste: Cacao Biological Station, 1030–1400 m: LACM 149071; Puntarenas: FincaZeledón, Río Coto, 1500 m: KU 129335; Finca Las Cruces 4.5 mi S San Vito de Java: LACM 114166, 121741; 1.6 km SSan Vito de Java, 1100 m: LACM 149073, 149076.

Anolis carpenteri—Costa Rica: Cartago: Turrialba, IICA Experimental Cacao Grove: MCZ 146765; Heredia: OTS Finca LaSelva: MCZ 174169; Finca La Suerte: SMF 81823; Limón: Finca La Lola: MCZ 128509–10; Tortuguero, next to Carib-bean Conservation Commission Camp: USNM 244860; San José: Caspirola: LSUMZ 52385. NICARAGUA: Matagalpa:12 km NE Matagalpa, 1100 m: KU 195056; Finca Tepeyac, 10.5 km N and 9 km E Matagalpa, 960 m: KU 85722; Río SanJuan: by Isla de Diamante on Río San Juan: OMNH 36370; Bartola, 10°58.37’N, 84°20.35’W, 30 m: SMF 80958, 81821–22. Panama: Bocas del Toro: south end of Isla Popa, 1 km E of Sumwood Channel: USNM 347200–04; Rambala: ZFMK52204; Laguna de Tierra Oscura, 3.7 km S of Tiger Key: USNM 348442; Colón: “Colón”: USNM 266152; Río Frijolesand Pipeline Road: USNM 266153.

Norps fortunensis—Panama: Chiriquí, Gualaca, Reserva Forestal Fortuna, 8º 43’04.7’’N, 82º13’42.1’’W, 1074 m: SMF86405–08; Reserva Forestal de Fortuna, along the Río Chiriquí, some 100 m from the mouth of Quebrada Bijau(8°45’11’’N, 82°11’07’’W), 1050–1075 m: MVUP 901; Reserva Forestal de Fortuna, along the Río Chiriquí, in front ofthe estación hidrometereológica del Instituto de Recursos Hidráulicos y Electrificación (IRHE) in Bijau, 1050–1075 m:



MVUP 756.Anolis fuscoauratus—Bolivia: Bení: Tributary of upper Río Bení: SMF 10967. Ecuador: Tena, 518 m: SMF 77252; near

Mera: SMF 77253; near Misahualli, 420 m: SMF 77254; Arutam, 1°47.28’S, 77°48.31’S, 760 m: SMF 78699. Panama:Darién: Río Chíco, near Avolinos: AMNH 57703. Venezuela: Iguapo, upper Orinoco: SMF 10971.

Anolis gruuo—Panama: Comarca Ngöbe Buglé: Nole Düima, Jadeberi, Serranía de Tabasará, from near the headwaters of RíoSan Félix, ca. 2 km N Escopeta Camp ca. 8°32’N, 81°50’W, 900 m: SMF 85416–19.

Anolis kemptoni—Panama: Chiriquí: Boquete, PNVB, 8º50’56’’N, 82º29’11’’W, 1610 m: SMF 85404–10; Bugaba, Bambito,Cerro La Pelota: SMF 85411–12, 85414–15; Parque Nacional La Amistad, 08º54’02,9’’N, 82º 36’38’’W, 1890 m: SMF85423–31; Parque Nacional La Amistad, 8°52’33.5’’N, 82°28’10.5’’W, 1700 m: MHCH 475–476, 497–502; headwatersof Río Chevo, 08º52’27,6’’N, 82º 44’31.7’’W, 1615 m: SMF 85448–50.

Anolis monteverde—Costa Rica: Puntarenas: 2 km E Santa Elena (10º20’30.5’N, 84º48’16.0’’W), 1550 m: SMF 86920, UCR20135–20136; road from Santa Elena to Reserva Santa Elena (10º19’54.8’N, 84º46’46.4’’W), 1575 m: SMF 85540; roadfrom Santa Elena to Reserva Santa Elena (10º20’29.6’N, 84º48’18.2’’W), 1560 m: SMF 85541–85548; Monteverde, 1545m: LACM 151313–151314; Monteverde, 1550 m: LACM 151315; 2.6 km ESE Monteverde, 1550–1580 m: LACM151316–151317; Monteverde, La Ventana, 1590 m: UCR 4971.

Anolis pseudokemptoni—Panama: Comarca Ngöbe Buglé: Nole Düima, Jadeberi, Serranía de Tabasará, La Nevera, 8°30’N,81°46’20’’W, 1600 m: SMF 85420–22.


koehler anolis tenorioensis

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