A new species of anole related to Anolis altae from Volcán Tenorio, Costa Rica (Reptilia, Squamata, Polychrotidae) Author Köhler, Gunther text Zootaxa 2011 3120 29 42 journal article 45849 10.5281/zenodo.205375 b838de53-aa90-4dcb-886d-2afdd7a11485 1175-5326 205375 Anolis tenorioensis sp. nov. Figs. 1 , 2 a, 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. fuscoauratus , A. gruuo , A. kemptoni , A. monteverde and A. pseudokemptoni in that it is short-legged (longest toe of adpressed 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 surfaces of body and limbs distinctly reticulated with dark brown pigment (versus immaculate dirty white or cream colored); and a male dewlap that is dark red with brown blotches (versus more or less uniformly reddish orange in A. altae and A. monteverde , uniform orange in A. carpenteri , posterior portion of male dewlap greenish beige, anterior portion reddish orange in A. fortunensis , uniform pinkish brown to red in A. fuscoauratus , more or less uniform dull orange in A. gruuo , posterior portion rose pink, anterior portion burnt orange, small cream colored basal blotch in some populations in A. kemptoni , posterior portion vinaceous, anterior portion orange, anterodorsal corner cream color in A. pseudokemptoni ); and by having a divided prenasal scale (versus usually a single elongated prenasal scale). It also differs from A. fortunensis , A. kemptoni , and A. monteverde by having a bilobate hemipenis (versus unilobate). Additionally, it can be differentiated from A. carpenteri by having a brownish dorsal ground color (versus greenish). Anolis tenorioensis is most similar to A. altae and A. monteverde , from which it further differs in several morphometric and pholidotic characteristics, such as relative head length (ratio head length / SVL) and number of 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 width 6.1 mm ; longest toe of adpressed hind limb reaching to tympanum; shank length 9.9 mm , shank length/head length ratio 0.93; longest finger of extended forelimb reaching nostril; longest finger of adpressed forelimb reaches a point 3.6 mm in front of anterior insertion of hind limbs. Most scales on snout keeled ( Fig. 4 ); 6 postrostrals; 7 scales between nasals; 2 prenasal scales, the lower one in contact with both rostral and first supralabial; scales in distinct prefrontal depression slightly wrinkled; supraorbital semicircles well developed, separated by a minimum of 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 overlapping superciliaries, the anterior one much larger than the posterior one; about 3 or 4 rows of small keeled scales extending between enlarged supraorbitals and superciliaries; parietal depression absent; interparietal scale well developed, 2.0 x 1.2 mm (length x width), surrounded by scales of moderate size; 3 scales present between interparietal and supraorbital semicircles; canthal ridge distinct, composed of 3 large (posterior two largest) and 5 small anterior 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 single row; 7 supralabials to level below center of eye; 5 suboculars broadly in contact with supralabials; ear opening 0.7 x 0.9 mm (length x height); mental distinctly wider than long, almost completely divided medially, bordered posteriorly 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 to level of axilla; dorsum of body with weakly keeled imbricate scales with rounded posterior margins, medial rows not enlarged, largest dorsal scales about 0.34 x 0.31 mm (length x width); about 47 medial dorsal scales in one head length; 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 0.29 mm (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 not enlarged, without whorls of enlarged scales, although an indistinct division in segments is discernible; a pair of slightly enlarged postcloacal scales present, about 0.60 mm wide; no tube-like axillary pocket present; scales on dorsal surface of forelimb keeled, imbricate, about 0.21 x 0.31 mm (length x width); digital pads dilated, dilated pad about 3 times width of non-dilated scales on distal phalanx; distal phalanx narrower than and raised from dilated pad; 24 (right)–26 (left) lamellae under phalanges II–IV of fourth toe; 8 scales under distal phalanx of fourth toe. 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. FIGURE 3. Holotype of Anolis tenorioensis (SMF 91985). SVL = 42.0 mm. Photos by Gunther Köhler. FIGURE 4. Head of holotype of Anolis tenorioensis (SMF 91985). Upper lateral view, middle dorsal view, lower ventral view of 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 1.0 mm. Photos by Gunther Köhler. 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 asulcate ridge 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 preservation. Photo by Gunther Köhler. The extended dewlap in life ( Fig. 2 a) has six horizontal gorgetal-sternal rows with a declining number of scales per row from base to tip (17–19 scales per row on basal portion, 5–9 towards tip); about 50 scales in marginal series. Coloration in life was recorded as follows: Dorsal surface of head Burnt Umber (22); dorsal ground color of body and limbs Grayish Horn Color (91), with shades of Buff (124) and some Dark Brownish Olive (129) pigmentation 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); dewlap Pratt’s Ruby (210) with Vandyke Brown (121) blotches. Coloration after 4 months preservation in 70% ethanol was recorded as follows: Dorsal ground color grayish brown with indistinct paler shadings, a cream colored temporal blotch, a dark brown interorbital bar, and faint crossbands on limbs and tail; ventral surfaces of head, body, limbs, and tail dirty pale grayish cream with a distinct dark brown reticulum; dewlap reddish (except damaged basal area) with dark brown and dirty white gorgetals, and with 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 ground color Raw Umber (123) with shades of Straw Yellow (56), a Mars Brown (223A) interorbital bar; snout Verona Brown (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 (specimens of Anolis altae and A. monteverde listed in Appendix I). Range is followed by mean value and one standard deviation in parentheses. For abbreviations see text.

A. altae 3 26 Ƥ 43	A. monteverde 3 8 Ƥ 8	A. tenorioensis 3 1 Ƥ 2
Maximum SVL 3 Ƥ	48.5 mm 50.2 mm	45.5 mm 47.0 mm	42.0 mm 45.0 mm
Tail length / SVL 3 Ƥ Tail diameter vertical / horizontal 3 Ƥ HL / SVL 3 Ƥ HL / HW 3 Ƥ IP / ear 3 Ƥ Shank length / SVL 3 Ƥ	1.89–2.30 (2.11±0.12) 1.81–2.22 (2.02±0.16) 1.04–1.64 (1.30±0.16) 0.95–1.40 (1.20±0.18) 0.27–0.29 (0.28±0.01) 0.25–0.28 (0.27±0.01) 1.56–1.82 (1.67±0.07) 1.50–1.78 (1.65±0.11) 1.26–6.86 (2.73±1.72) 0.82–5.81 (2.73±1.98) 0.20–0.25 (0.23±0.01) 0.18–0.25 (0.21±0.03)	1.81–1.93 (1.81±0.05) 1.65–1.93 (1.78±0.12) 1.05–1.43 (1.22±0.11) 1.11–1.31 (1.24±0.09) 0.27–0.29 (0.28±0.01) 0.25–0.28 (0.27±0.01) 1.63–1.74 (1.69±0.04) 1.60–1.75 (1.69±0.07) 1.71–3.67 (2.61±0.60) 2.01–4.40 (2.95±1.05) 0.22–0.24 (0.22±0.08) 0.20–0.22 (0.21±0.01)	— 1.64–1.69 1.00 1.13–1.33 0.25 0.24–0.25 1.75 1.70–1.82 3.81 1.60–4.00 0.24 0.21–0.22
Axilla–groin distance / SVL 3 Ƥ	0.34–0.46 (0.42±0.03) 0.40–0.52 (0.46±0.05)	0.38–0.46 (0.43±0.03) 0.40–0.49 (0.44±0.03)	0.45 0.38–0.47
DHL 3 Ƥ	44–74 (57.83±7.71) 40–66 (50.72±10.60)	49–78 (62.13±9.54) 49–68 (55.5±8.44)	47 47–48
VHL 3 Ƥ	34–56 (43.52±5.53) 24–48 (34.70±9.25)	47–61 (54.75±4.43) 40–52 (46.75±5.34)	48 43–46
SAM 3	122–150 (136.33±7.54)	140–156 (148.25±5.90)	101
Ƥ	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 Number of scales between IP and SS Number of scales between SO and SPL	0–3 (1.46+±0.70) 1–4 (2.13±0.87) 0	1–3 (1.78±0.48) 2–3 (2.33±0.45) 0	1–2 2–3 0
Number of SPL to level below center of eye Number of INL to level below center of eye Total number of loreals Number of horizontal loreal scale rows Number of postrostrals Number of postmentals	5–8 (6.80±0.68) 5–8 (6.43±0.79) 24–58 (41.73±8.55) 4–8 (6.07±0.86) 4–8 (6.27±0.78) 4–7 (5.60±0.74)	7–9 (7.50±0.63) 7–8 (7.63±0.50) 37–68 (47.81±8.41) 6–9 (6.63±0.81) 5–9 (7.06±1.06) 4–6 (5.63±0.81)	6–7 6–7 27–28 5 6 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 Etymology. The name tenorioensis is derived from Volcán Tenorio where the type series of the species was collected and to where it is probably restricted, and the Latin suffix - ensis (denoting place or locality). FIGURE 7. View of the cloud forest at Volcán Tenorio (taken at a point about 1000 m elevation), Costa Rica. Photo by Donald Varela 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 vegetation (mostly on fern leaves) within the forest (Premontane Rainforest fide Savage 2002 ) near an open area that is flooded most of the year and known locally as Laguna Danta (see also Figs. 7 and 8 ). At the type locality, a recently fallen 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 highlands of Volcán Tenorio in the Cordillera de Guanacaste, Costa Rica . It occurs in a protected area (Parque Nacional Volcán Tenorio). Only three specimens of this species are known to science. However, given its presumably small geographic range, it seems justified to classify A. tenorioensis as Critically Endangered based on criterion B.1.a. of the IUCN Red List Categories and Criteria (IUCN 2011).