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The Anolis heterodermus group comprises eight big-headed and short-legged lizard species from the highlands of the northernmost South American Andes. Recent studies revealed unknown lineages within this group that had previously been categorized as a species complex. By widely sampling and applying an integrative taxonomic framework, we (1) assessed the species diversity of the group using a molecular dataset (two mitochondrial and one nuclear markers) along with an inclusive morphological study (scalation, scale configuration and ornamentation, morphometrics, and dewlap and body colour patterns); and (2) we inferred the evolutionary relationships within this species group. Our analyses confirmed the formerly reported differentiation between populations of those high-altitude lizards, and we identified several unknown evolutionary lineages. Our results provided evidence for the existence of nine distinct, independently evolving evolutionary lineages in the heterodermus group. As a result, we described two morphologically and genetically highly distinct lineages as species new to science (A. quimbayasp. nov. and A. tequendamasp. nov.). We redescribed A. heterodermus and erected as a valid species Anolis richteri, a previously described synonym of A. heterodermus. A taxonomic key for the identification of species of the Phenacosaurus clade was presented. The identification of two additional poorly-known lineages suggested that the diversity of this group of lizards is still unknown; therefore, it is necessary to establish measurements for the group´s conservation, as well as to perform fieldwork and revision of herpetological collections to identify possible hidden diversity within the group.

Anolis riparius, new species Holotype. UCR 5579 (Fig 6) collected on 7 September 2009 by Douglas Robinson and his students; an adult male from Pacuarito river, Pacuarito 10 Km S-SW of Pérez Zeledón, 9.31390 N, - 83.77220 W, 880 masl, San José Province, Costa Rica. Paratypes (all from Costa Rica: San José Province). Adult males: SMF 82183–82184 (collected 9 October 2002 by Axel Fläschendräger) from road between Platanillo and San Isidro, 9.31056 N; - 83.77500 W, ca. 850 masl; SMF 92428 (collected 15 February 2010 by Gunther Köhler) from 1 km N Cedral 9.35789 N, - 83.56094 W, 1425 masl; UCR 7893 (collected 24 August 1979 by Douglas Robinson) from I.85 Km NE Alfombra, 9.3167 N, - 83.7722 W, ca. 930 masl; MCZ-R 18171 (collected 15 December 2007 by Steve Poe and Mason J. Ryan) from Alfombra, 12 Km S-SW of Pérez Zeledón (9.31670 N, - 83.77220 W, 930 masl); MSB 95201 (collected on 10 January 2009 by Mason J Ryan and Ian Latella) from Alfombra, 12 Km S-SW of Pérez Zeledón (9.31670 N, - 83.77220 W, 930 masl). Adult females: SMF 82185–82186 (collected 9 October 2002 by Axel Fläschendräger) from road between Platanillo and San Isidro (ca. 850 masl); SMF 83093–83094 (collected 9 October 2002 by Axel Fläschendräger) from near Platanillo (ca. 850 masl); SMF 92429 (collected 15 February 2010 by Gunther Köhler) from 1 km N Cedral (9.35789 N, - 83,56094 W, 1425 masl); UCR 5580 (same data of the holotype); UCR 15959 (collected 3 March 2001 by Mason J Ryan) from Mollejones (9.22930 N, - 83.64200 W, 800 masl); UCR 16162 (collected 2 July 2001 by Mason J Ryan) from Alfombra, 12 Km S-SW of Pérez Zeledón (9.31670 N, - 83.77220 W, 930 masl); MCZ R-186143 (collected 15 December 2007 by Steve Poe and Mason J Ryan) from Alfombra, 12 Km S-SW of Pérez Zeledón (9.31670 N, - 83.77220 W, 930 masl); MSB 95200 (collected 15 December 2007 by Steve Poe and Mason J Ryan) from Alfombra, 12 Km S-SW of Pérez Zeledón (9.31670 N, - 83.77220 W, 930 masl); UCR 16638 (collected 15 March 2002 by Franklin Aguilar) from Fila Piedras Blancas area (9.50130 N, - 83.91280 W, 900 masl) in the Zona Protectora Los Santos in the Río Savegre basin. Diagnosis. Anolis riparius sp. nov. and A. aquaticus are the only semiaquatic Middle American anoles that have a large orange-red dewlap with yellow in males (Fig. 6B). Anolis riparius sp. nov. differs from A. aquaticus in possessing smaller dorsal scales of the head and body (e.g., 15–19 scales across the snout between the second canthals in A. riparius sp. nov. versus 7–14 in A. aquaticus, Fig. 5B), and is phylogenetically distinct (see below). Anolis riparius sp. nov. is distinguished from the other Middle American semiaquatic anoles as follows: from A. robinsoni sp. nov. by male dewlap color (chocolate brown with ill-defined brick red horizontal stripes in A. robinsoni sp. nov.); from A. lionotus and A. poecilopus by male dewlap color (solid yellow-orange in A. lionotus and A. poecilopus), lack of postcloacal scales in males (usually present in A. lionotus and A. poecilopus), and possession of zero to three enlarged rows of middorsal scales (a broadband of 10–24 enlarged middorsal scale rows in A. poecilopus and A. lionotus); from A. barkeri by smaller size (maximum SVL 73 mm in A. riparius sp. nov., 91 mm in A. barkeri), presence of distinctly expanded toe pads (ratio width of expanded pads / width of distal phalanx 1.8–2.2 in A. riparius sp. nov. versus 1.4–1.6 in A. barkeri), and presence of a single row of middorsal caudal scales (double row in A. barkeri). Description of the holotype (Figure 6). Adult male as indicated by dewlap and everted hemipenes SVL 73.3 mm; HL 20.1 mm; HL/SVL 0.27; HW 10.7 mm; EOH 1.64 mm; IL 1.45 mm; IL/EOH.88; SLS-I 0.1; AGL 31.26 mm; FL 18.34 mm; T4L 15.95 mm; T4W 1.54 mm; TL 135.00 mm, tail complete; TL/SVL 1.84; TH 5.994 mm and TW 3.42 mm. 18 scales across the snout between second canthals; two scales separate the nasal opening from rostral scale; ten postrostral scales between supralabials; 12 supralabial scales from rostral scale to level of middle of eye; four scales separating supraorbital semicircles; 2 rows of scales separate subocular scales from supralabials; one slightly elongate superciliary scale; nine scales separating interparietal scale from supraorbital semicircles; four enlarged scales in supraocular disk; eight loreal scales in column just anterior to eye; 12 postmental scales posteriorly in contact with mental scale between infralabials; 15 scales in the loreal region; 13 rows of single scales in the dewlap (21–48 scales per row, rows somewhat irregular); seven enlarge middorsal scale row; 16 dorsal scales in 5% of SVL; 12 ventral scales in 5% of SVL; 220 scales around midbody; 15 expanded lamellae under fourth toe; 23 small middorsal caudal scales before the first large caudal scale (Fig. 6D). Preoccipital, enlarged infralabial, tail crest, middorsal, postcloacal scales absent; supraocular disc presents with small scales. Frontal region of head concave; scales in supraocular disc unicarinate; rostral scale with very weak cleft and overlaps mental scale; ear opening vertically ovoid; dorsal edge of ear lacks ornamentation; mental scale completely divided posteriorly and with its posterior edge convex; dewlap extends well on to chest with anterior insertion at level of anterior portion of the eye and without protruding scales at distal edge; dorsal scales strongly keeled and uniform in size; ventral scales in diagonal rows keeled and larger than dorsal scales; lateral scales homogeneous; tail is compressed and triangular in cross-section with the base taller than wide; middorsal caudal scales in single row and its size is less than three times than the adjacent scales (Fig. 6D); the longest toe of adpressed limb reaches anterior to eye and, supradigital scales keeled and multicarinate. Color in Life (based on field notes and color photographs; Fig. 6A and 6B). Dewlap yellow with orange-red horizontal streaks. Dorsal ground color olive to chestnut brown with transverse olive-green bars across dorsum down flanks extending to tail. A dirty cream-colored line extends posteriorly from the labial scales down the side of the body. Scattered small greenish spots are present on the dorsum. The dorsum and tail are marked with olive-green transverse bars. The limbs are marked with pale green to yellowish transverse bars with small punctuations. Variation. Size of reproductive males (69.6 ± 5.3, 65.8–73.3 mm) was larger than females (58.0 ± 6.0, 49.1–65.1 mm). Table 1 shows variation in the lepidosis characters of Anolis riparius sp. nov.. Individuals exhibited little variation in coloration. Although some individuals were dark, the amount and distribution of the light bands and dots were similar among all specimens. Hemipenis morphology. The completely everted hemipenis of SMF 92428 (Fig. 7B) is a stout bilobed organ. The sulcus spermaticus is bordered by well-developed sulcal lips and opens into two broad concave areas, one on each lobe. A small asulcate ridge is present. The apex is strongly calyculate, truncus with transverse folds. Etymology. The specific epithet is an adjective taken directly from Latin (riparius = of stream banks) about the riparian habitat of this species. Distribution. Anolis riparius sp. nov. is known from streams in the riparian gallery forests of the Pacuar, Guabo, and Savegre river basins (Fig. 1) from 100 to 1450 masl. This species inhabits lowland and premontane tropical wet forest life zones. Ecology. Little is known about the ecology and life history of Anolis riparius sp. nov. but it seems to share lifehistory traits with A. aquaticus and A. robinsoni sp. nov.. What follows is based mainly on field observations by MJR from Alfombra and additional unnamed streams of the Rio Guabo watershed. This species may be common in small order and headwater streams but absent from some other streams within a watershed. During the day this species may be found perched on rocks and boulders along streams and can jump into the water or crawl into interstitial cracks between rocks and boulders when pursued. At night these lizards are found sleeping on moss mats, fern fronds, and other large leafy riparian vegetation, especially near the splash zone around boulders. We have also found juveniles and adults sleeping on vertical rock walls and boulders in and next to waterfalls. We have observed these lizards sleeping from 0.5 to 2.0 meters above the ground. At Alfombra on 21 December 2007 MJR and SP found a congregation of seven gravid females on the same fern-covered boulder. One meter from the sleeping females was a cluster of 6 loosely aggregated eggs that were stuck to the moss on the same large boulder. Morphological comparisons of the two new species and Anolis aquaticus . Anolis aquaticus is the smallest species in this complex. It has fewer scales for all traits except for the number of scales between the suboculars and supralabials (Table 1 ). Anolis aquaticus has relatively larger dorsal scales, whereas A. robinsoni sp. nov. and A. riparius sp. nov. have small, granular dorsal scales that result in a higher number of total of scales around the body (more than 160, whereas A. aquaticus has 160 or fewer). The number of scales between the second canthals is close to half in number in A. aquaticus relative to the other species (Table 1, Fig. 4B, 5B, 6B), indicating the larger size of the scales in A. aquaticus. The number of small middorsal caudal scale varies between 0 to 10 in A. aquaticus (Fig. 5C) while the other species has more than 20 small middorsal caudal scales (Fig. 4D and 4D). However, the most striking feature separating one of the three species of this group is the color of the male dewlap. The red and yellow dewlap of A. aquaticus and A. riparius sp. nov. (Fig. 5A and 6B) contrasts with the dark brown dewlap of A. robinsoni sp. nov. (Fig. 4B). We did not find any difference in hemipenes morphology among A. aquaticus, A. riparius sp. nov., and A. robinsoni sp. nov. (Fig. 7).

Anolis robinsoni, new species Holotype. UCR 2463 (Fig. 4) collected 15 March 1969 by Oscar Blanco, an adult male from Palma stream bridge 5.1 km south of Santa Marta de Santiago de Puriscal, 9.79230 N, - 84.39530 W, ca. 800 masl, San José Province, Costa Rica. Paratypes ( all from Costa Rica: San José Province ). Adult males: SMF 92437 (collected 27 March 2010 by Gunther Köhler) from Zona Protectora El Rodeo (Universidad para la Paz), 9.90392 N, - 84.28169 W, ca. 825 masl; SMF 92438 (collected 27 March 2010 by Gunther Köhler) from Zona Protectora El Rodeo (Universidad para la Paz), 9.90355 N, - 84.28192 W, ca. 780 masl; UCR 2464 (same data of the holotype); UCR 16048 (collected 28 October 2001 by Mason J Ryan, Robert Puschendorf and Brian Kubicki) from the west slope of the Cerros de Escazú, Río Jaris (9.88460 N, - 84.27650 W, 500 ca. masl), close to the Zona Protectora El Rodeo (Universidad para la Paz); MCZ R-186162 (collected 27 December 2007 by Steve Poe and Mason J. Ryan) from Quebrada La Palma in Alto Palma de Puriscal (9.78890 N, - 84.39440 W, ca. 1000 masl). Adult females: SMF 92439, 92450–51 (collected 27 March 2010 by Gunther Köhler) from Zona Protectora El Rodeo (Universidad para la Paz), 9.90364 N, - 84.28182 W, ca. 820 masl; UCR 2558 (collected 15 March 1969 by Oscar Blanco and Douglas Robinson) and MCZ R-186161 (collected 27 December 2007 by Steve Poe and Mason J. Ryan) both of them from Quebrada La Palma in Alto Palma de Puriscal (9.78890 N, - 84.39440 W, ca 1000 masl). Diagnosis. Anolis robinsoni sp. nov. is a semiaquatic anole that is distinguished from all other Anolis of Central America by male dewlap coloration (chocolate brown with ill-defined brick red horizontal streaks, Fig. 4B). The only other species with which A. robinsoni sp. nov. might be confused in the field are the other Middle American semiaquatic anoles A. aquaticus, A. lionotus, A. poecilopus, and A. barkeri. Anolis lionotus and A. poecilopus are easily distinguished from A. robinsoni sp. nov. by male dewlap color (dull orange-yellow in these species, chocolate brown with ill-defined brick red horizontal streaks in A. robinsoni sp. nov.), presence of enlarged postcloacal scales in most males (absent in A. robinsoni sp. nov.), and presence of a longitudinal zone of 10–24 enlarged middorsal scales (0–3 scales enlarged in A. robinsoni sp. nov.). Anolis aquaticus may be distinguished from A. robinsoni sp. nov. by male dewlap color (orange-red with yellow in A. aquaticus, Fig. 5A) and possession of larger dorsal scales of the head and body (e.g., 15–20 scales across the snout between second canthals in A. robinsoni sp. nov. (Fig. 4C), 7–14 in A. aquaticus; Fig. 5B, Table 1) and the size of the middorsal caudal row scales relative to adjacent scales (no more than two times in A. robinsoni sp. nov. (Fig. 4D), more than three times in A. aquaticus (Fig 5C). Anolis barkeri is most easily distinguished from A. robinsoni sp. nov. by its larger size (maximum SVL 91 mm in A. barkeri, 74 mm in A. robinsoni sp. nov.), lack of distinctly expanded toe pads (ratio width of expanded pads/width of distal phalanx 1.8–2.2 in A. robinsoni sp. nov. versus 1.4–1.6 in A. barkeri), and presence of a double row of middorsal caudal scales (single row in A. robinsoni sp. nov.). Description of the holotype (Fig. 4). Adult male as indicated by dewlap and everted hemipenes; SVL 63.9 mm; HL 17.5 mm; HL/SVL 0.28; HW 10.1 mm; EOH 2.36 mm; IL 1.18 mm; IL/EOH 0.50; SLS-I 0.18 mm mm; AGL 23.1 mm; FL 16.6 mm; T4L 15.1 mm; T4W 1.42 mm; TL 10.4 mm, tail complete; TL/SVL 0.38; T4L 15.1 mm and T4W 1.42 mm. 18 scales across the snout between second canthals; two scales separate the nasal opening from rostral scale; nine postrostral scales between supralabials; nine supralabial scales from rostral scale to level of middle of eye; six scales separating supraorbital semicircles; one rows of scales separate subocular scales from supralabials; two slightly elongate superciliary scale; eight scales separating interparietal scale from supraorbital semicircles; 12 enlarged scales in supraocular disk; 11 loreal scales in column just anterior to eye; 11 postmental scales posteriorly in contact with mental scale between infralabials; 16 scales in the loreal region; 19 rows of single scales in the dewlap (10–42 scales per row, rows somewhat irregular); 2 middorsal scale rows slightly larger than adjacent scales; 16 dorsal scales in 5% of SVL; 12 ventral scales in 5% of SVL; 220 scales around midbody; 23 expanded lamellae under fourth toe; 26 small middorsal caudal scales before the first large caudal scale (Fig. 4D). Preoccipital, enlarged infralabial, tail crest, middorsal, and postcloacal scales absent; supraocular disc presents small scales. Frontal region of head concave; scales in supraocular disc unicarinate; rostral scale with weak cleft and overlaps mental scale; ear opening vertically ovoid; dorsal edge of ear lacks ornamentation; mental scale partially divided posteriorly and with its posterior edge convex; dewlap extends well on to chest with anterior insertion at level of anterior portion of the eye and without protruding scales at distal edge; dorsal scales small, keeled with two middorsal scale rows slightly larger than adjacent scales; ventral scales in diagonal rows, keeled and larger than dorsal scales; lateral scales homogeneous; tail is compressed and triangular in cross-section with the base taller than wide; middorsal caudal scales in single row and its size is less than three times than the adjacent scales (Fig. 4D); the longest toe of adpressed limb reaches anterior to eye and, supradigital scales keeled and multicarinate. Color in life (male based on field notes and color photographs; Fig. 4A and 4B). The dewlap of males in life is chocolate brown with lighter orange-brown bars located centrally. The dorsal ground color is olive to chestnut brown with transverse olive–green bars across dorsum down flanks extending to tail. The head is uniformly dark or light brown. A cream-colored line extends posteriorly from the labial scales down the side of the body. Scattered small greenish spots are present on the dorsum. Scales on the tip of the snout with black pigmentation. The dorsum and tail are marked with olive–green transverse bars and the limbs marked with pale green to yellowish transverse bars with small punctuations. Variation. Anolis robinsoni sp. nov. showed variation in the SVL of reproductive males (58.6 ± 11.0, 42.1–73.8 mm). Table 1 shows the variation in lepidosis. The gular region of females is white with brown streaks. Females have a small brown to black dewlap. Transverse, dorsal olive-green bands of males varied between 7–8, with 10–12 tail bands; females had 7–8 olive–green dorsal transverse bars with 9–11 tail bands. In life, the dorsal coloration of both sexes may vary with transverse bands darker when cold or stressed (e.g., when held in hand). Hemipenis morphology. The almost completely everted hemipenis of SMF 92438 (Fig. 7A) is a stout bilobed organ. The sulcus spermaticus is bordered by well-developed sulcal lips opening into two broad concave areas, one on each lobe. A small asulcate ridge is present. The apex is strongly calyculate, truncus with transverse folds. Etymology. The specific epithet is a patronym in honor of the late Douglas C. Robinson, who contributed enormously to the herpetological knowledge of Costa Rica through extensive field collections and by inspiring a generation of Costa Rican biologists. He received his Ph.D. from Texas A&M University in 1968 and studied the Mexican semiaquatic anole, Anolis barkeri. Distribution. Anolis robinsoni sp. nov. is known from the riparian gallery forests of the Río Candelaria basin (Fig. 1), the foothills of the Montañas de Turrubares in the northwestern section of the Cordillera de Talamanca, and Río Jaris along the western border of the Zona Protectora El Rodeo. These areas are within the Tropical Wet and Humid Premontane forest life zones (Holdridge 1967). Anolis robinsoni sp. nov. has been found at elevations between 500 and 1100 masl. Ecology. Most of the following observations are from Márquez–Baltán (1994) and Márquez et al. (2005) from Quebrada La Palma. During the day this species is found on bare rocks and boulders along streams with dense canopy cover. At night it is found sleeping on moss and vegetation on boulders near the splash zones of small and medium waterfalls. Both sexes and juveniles were found no more than two meters from the edge of a stream. The operational sex ratio is approximately equal proportions of males to females (Márquez et al. 2005), and males share territories with one to three females. The estimated population density varied from 100 to 200 individuals per hectare and population size fluctuated monthly with a peak during the dry season. Reproduction occurs throughout the year with a lull in the wet season (May to November). Females lay 1 egg per clutch in cracks and interstitial spaces of boulders in streams. Up to 6 eggs have been found together suggesting that this species lays eggs in communal nest sites. Incubation times ranged between 75–82 days, and hatchlings are approximately 25 mm SVL (Márquez et al. 2005).

Chaves, Gerardo, Ryan, Mason J., Bolaños, Federico, Márquez, Cruz, Köhler, Gunther, Poe, Steven (2023): Two new species of semiaquatic Anolis (Squamata: Dactyloidae) from Costa Rica. Zootaxa 5319 (2): 249-262, DOI: 10.11646/zootaxa.5319.2.6, URL: http://dx.doi.org/10.11646/zootaxa.5319.2.6

Anolis sagrei Duméril & Bibron, 1837 — Non-native; Established. Agama Sagrei Duméril & Bibron, 1837: 149–152. Syntypes (7): MNHN 2430, 2430A, 2430B, 6795, 6795A 6797, 6797A, by original designation (see Brygoo [1989: 90]); MCZ 2172 also considered a syntype (see McCranie & Ģnther [2015: 166]) based on Underwood & Stuart (1959: 17) who noted “? Cotype M.C.Z. 2172, received from the Jardin des Plantes” (see Brygoo [1989: 90]). Type locality: “habite l’île de Cuba ” (= Cuba), restricted to “the city of La Habana, Habana ” (= Havana, La Habana Province, Cuba) by Ruibal (1964: 490). Brown Anole (Figure 11D) Singapore records. Norops sagrei —H.H. Tan & K.K.P. Lim, 2012: 359–362 (Gardens by the Bay).—K.W.C. Leong & K.K.P. Lim, 2015: 123, 124 (Marina Bay South Gardens).—Woo & Wee, 2015: 61 (Clover Rise [Bishan]).—H.H. Tan & Zhou, 2016: 171–172 (Gardens by the Bay).—C.L.Y. Tan et al., 2017: 48 (Singapore Botanic Gardens).—A. Tay, 2019c: 82 (Thomson Road).—H. Cai & Z. Wu, 2020: 147 (Jurong Lake Gardens).—A. Tay & J. Lim, 2021: 1 (Bah Soon Pah Road; Thomson Road; Toa Payoh Central).— E.K. Chua, 2022: 88 (Satay by the Bay [= Gardens by the Bay). Anolis sagrei —E. S. C. Goh & R.W.X Seah. 2020: 190 (Kovan area, Tampines Road [compound of Horizon Floral & Design Pte Ltd]).—J. Chua, 2021: 1 (Marina Bay).—N.T.-L. Lim, 2022: 1 (Keat Hong Close).—A. Tay, 2022: 1 (Seletar West Farmway 7). Remarks. Anolis sagrei is a recently introduced and established species being first brought in as stowaways with ornamental plants at Gardens by the Bay in 2012 (Tan & Lim 2012). Native to the Bahamas, Swan Island, and Cuba (Nicholson et al. 2012), A. sagrei has been introduced to at least nine countries (Global Invasive Species Database 2019a).At the time of discovery, numerous adult and hatchling individuals were recorded, and courtship behaviour was observed (Tan & Lim 2012). Since the ornamental plants were in place by October 2011, A. sagrei was likely already reproducing and increasing in numbers, especially since it has a short breeding cycle capable of causing a marked surge in population size (Norval et al. 2012). Report of a single individual in Bishan, a locality 15 km north of Gardens by the Bay on 9 March 2015 (Woo & Wee 2015), SBG on 14 April 2017 (Tan et al. 2017), Hort Park in July 2017 (ZRC.2.7280), Toa Payoh on 3 June 2021 (Tay & Lim 2021), and Choa Chu Kang on 30 April 2022 (Lim 2022) suggests that A. sagrei may be expanding its range throughout Singapore; however, only a single individual was observed at each location. Furthermore, individuals photographed at plant nurseries along Thomson Road on 23 May 2019 and 5 March 2021 (Tay 2019c; Tay & Lim 2021), Bah Soon Pah Road on 6 February 2021 (Tay & Lim 2021), and Seletar West Farmway 7 on 12 December 2022 (Tay 2022) hint that more stowaways are being introduced. This indeed may be the case as an established population has now been detected at JLG (Cai & Wu 2020), which underwent renovation in 2018 and 2019 that included wide-scale planting, and one individual was photographed at a nursery in Kovan on 28 October 2020 (Goh & Seah 2020). In other parts of the world where A. sagrei was introduced, it is considered an invasive species because it outcompetes competitors and greatly decimates arthropod populations (Huang 2008; Meshaka 2011). At first it was believed that A. sagrei’s impact on native species would be minimal as it was restricted to Gardens by the Bay, which is built on reclaimed land. However, if A. sagrei is indeed expanding its range throughout Singapore, then it may soon begin competing with native species. Occurrence. Established populations are restricted to Gardens by the Bay, Marina Bay South, and Jurong Lake Gardens, with a few isolated records from elsewhere. Common. Singapore conservation status. Not Applicable. Conservation priority. None, non-native species. IUCN conservation status. Least Concern [2020]. LKCNHM & NHMUK Museum specimens. Hort Park: ZRC.2.7280 (Jul-2017); Marina Bay South Gardens: ZRC.2.6988 (18-Oct-2012), ZRC.2.7024 (31-May-2013). Additional Singapore museum specimens. Singapore (no locality): No specimens. Singapore localities. Bishan—Bah Soon Pah Road—Gardens by the Bay—Hort Park—Keat Hong Close— Kovan—Jurong Lake Gardens—Marina Bay South—Seletar West Farmway 7—Singapore Botanic Gardens—Thomson Road—Toa Payoh.

Figueroa, Alex, Low, Martyn E.Y., Lim, Kelvin K.P. (2023): Singapore's herpetofauna: updated and annotated checklist, history, conservation, and distribution. Zootaxa 5287 (1): 1-378, DOI: https://doi.org/10.11646/zootaxa.5287.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5287.1.1

Anolis sagrei Duméril & Bibron, 1837 — Non-native; Established. Agama Sagrei Duméril & Bibron, 1837: 149–152. Syntypes (7): MNHN 2430, 2430A, 2430B, 6795, 6795A 6797, 6797A, by original designation (see Brygoo [1989: 90]); MCZ 2172 also considered a syntype (see McCranie & Ģnther [2015: 166]) based on Underwood & Stuart (1959: 17) who noted “? Cotype M.C.Z. 2172, received from the Jardin des Plantes” (see Brygoo [1989: 90]). Type locality: “habite l’île de Cuba ” (= Cuba), restricted to “the city of La Habana, Habana ” (= Havana, La Habana Province, Cuba) by Ruibal (1964: 490). Brown Anole (Figure 11D) Singapore records. Norops sagrei —H.H. Tan & K.K.P. Lim, 2012: 359–362 (Gardens by the Bay).—K.W.C. Leong & K.K.P. Lim, 2015: 123, 124 (Marina Bay South Gardens).—Woo & Wee, 2015: 61 (Clover Rise [Bishan]).—H.H. Tan & Zhou, 2016: 171–172 (Gardens by the Bay).—C.L.Y. Tan et al., 2017: 48 (Singapore Botanic Gardens).—A. Tay, 2019c: 82 (Thomson Road).—H. Cai & Z. Wu, 2020: 147 (Jurong Lake Gardens).—A. Tay & J. Lim, 2021: 1 (Bah Soon Pah Road; Thomson Road; Toa Payoh Central).— E.K. Chua, 2022: 88 (Satay by the Bay [= Gardens by the Bay). Anolis sagrei —E. S. C. Goh & R.W.X Seah. 2020: 190 (Kovan area, Tampines Road [compound of Horizon Floral & Design Pte Ltd]).—J. Chua, 2021: 1 (Marina Bay).—N.T.-L. Lim, 2022: 1 (Keat Hong Close).—A. Tay, 2022: 1 (Seletar West Farmway 7). Remarks. Anolis sagrei is a recently introduced and established species being first brought in as stowaways with ornamental plants at Gardens by the Bay in 2012 (Tan & Lim 2012). Native to the Bahamas, Swan Island, and Cuba (Nicholson et al. 2012), A. sagrei has been introduced to at least nine countries (Global Invasive Species Database 2019a).At the time of discovery, numerous adult and hatchling individuals were recorded, and courtship behaviour was observed (Tan & Lim 2012). Since the ornamental plants were in place by October 2011, A. sagrei was likely already reproducing and increasing in numbers, especially since it has a short breeding cycle capable of causing a marked surge in population size (Norval et al. 2012). Report of a single individual in Bishan, a locality 15 km north of Gardens by the Bay on 9 March 2015 (Woo & Wee 2015), SBG on 14 April 2017 (Tan et al. 2017), Hort Park in July 2017 (ZRC.2.7280), Toa Payoh on 3 June 2021 (Tay & Lim 2021), and Choa Chu Kang on 30 April 2022 (Lim 2022) suggests that A. sagrei may be expanding its range throughout Singapore; however, only a single individual was observed at each location. Furthermore, individuals photographed at plant nurseries along Thomson Road on 23 May 2019 and 5 March 2021 (Tay 2019c; Tay & Lim 2021), Bah Soon Pah Road on 6 February 2021 (Tay & Lim 2021), and Seletar West Farmway 7 on 12 December 2022 (Tay 2022) hint that more stowaways are being introduced. This indeed may be the case as an established population has now been detected at JLG (Cai & Wu 2020), which underwent renovation in 2018 and 2019 that included wide-scale planting, and one individual was photographed at a nursery in Kovan on 28 October 2020 (Goh & Seah 2020). In other parts of the world where A. sagrei was introduced, it is considered an invasive species because it outcompetes competitors and greatly decimates arthropod populations (Huang 2008; Meshaka 2011). At first it was believed that A. sagrei’s impact on native species would be minimal as it was restricted to Gardens by the Bay, which is built on reclaimed land. However, if A. sagrei is indeed expanding its range throughout Singapore, then it may soon begin competing with native species. Occurrence. Established populations are restricted to Gardens by the Bay, Marina Bay South, and Jurong Lake Gardens, with a few isolated records from elsewhere. Common. Singapore conservation status. Not Applicable. Conservation priority. None, non-native species. IUCN conservation status. Least Concern [2020]. LKCNHM & NHMUK Museum specimens. Hort Park: ZRC.2.7280 (Jul-2017); Marina Bay South Gardens: ZRC.2.6988 (18-Oct-2012), ZRC.2.7024 (31-May-2013). Additional Singapore museum specimens. Singapore (no locality): No specimens. Singapore localities. Bishan—Bah Soon Pah Road—Gardens by the Bay—Hort Park—Keat Hong Close— Kovan—Jurong Lake Gardens—Marina Bay South—Seletar West Farmway 7—Singapore Botanic Gardens—Thomson Road—Toa Payoh., Published as part of Figueroa, Alex, Low, Martyn E. Y. & Lim, Kelvin K. P., 2023, Singapore's herpetofauna: updated and annotated checklist, history, conservation, and distribution, pp. 1-378 in Zootaxa 5287 (1) on pages 96-97, DOI: 10.11646/zootaxa.5287.1.1, http://zenodo.org/record/7960319, {"references":["Brygoo, E. R. (1989) Les types d'Iguanides (Reptiles, Sauriens) du Museum national d'Histoire naturelle Catalogue. Bulletin du Museum nationale d'Histoire naturelle, Paris, Series 4 A, 11 (3, Supplement), 1 - 112. https: // doi. org / 10.5962 / p. 288302","Chua, J. (2021) Biodiversity Record: Golden gliding snakes at Marina Bay. Nature in Singapore, 14 (e 2021120), 1 - 2.","Lim, N. T. - L. (2022) Biodiversity Record: A brown anole at Choa Chu Kang. Nature in Singapore, 15, e 2022047.","Global Invasive Species Database. (2019 a) Species profile: Norops sagrei. Available from: http: // www. iucngisd. org / gisd / speciesname / Norops + sagrei (accessed 14 January 2021)","Cai, H. & Wu, Z. (2020) Brown anoles at Jurong Lake Gardens. Singapore Biodiversity Records, 2020, 147 - 148.","Goh, E. S. C. & Seah, R. W. X. (2020) Observation of a brown anole at the Kovan area. Singapore Biodiversity Records, 2020, 190.","Huang, S. - C., Norval, G. & Tso, I. - M. (2008) Predation by an exotic lizard, Anolis sagrei, alters the ant community structure in betelnut palm plantations in southern Taiwan. Ecological Entomology, 33 (5), 569 - 576. https: // doi. org / 10.1111 / j. 1365 - 2311.2008.00995. x","Meshaka, Jr., W. E. (2011) A runaway train in the making: The exotic amphibians, reptiles, turtles, and crocodilians of Florida. Herpetological Conservation & Biology, 6, 1 - 101."]}

Cutaneous sensory organs are characteristic of many squamate lineages. Such organs may occur on the surface of scales as button-like, circular protuberances set off from their surroundings by a noticeable boundary, often taking the form of a moat or furrow. They may be relatively unadorned, clad with the surface micro-ornamentation of the scales on which they are carried, or they may carry one or more bristles of varying length and surface ornamentation. Such bristles may extend away from the body of the organ to interface with the surrounding environment or to contact adjacent scales. Cutaneous sensory organs have been physiologically demonstrated to have a mechanoreceptive function but have also been posited to potentially be involved with additional sensory modalities. Their distribution and structure across the body surface has been shown to be unequal, with some regions being much more extensively endowed than others, indicative of regional differential sensitivity. The digits of Anolis (Iguania: Dactyloidae) carry adhesive toepads that are convergent with those of geckos (Gekkota). Geckos exhibit a high density of cutaneous sensory organs on their toepads and their form and distribution has been associated with the operation and control of the toepads during locomotion. Investigation of the form and topographical distribution of cutaneous sensory organs on the toepads of Anolis shows them to be convergent in these attributes with those of geckos and quite distinct from those of the ancestrally padless Iguana (Iguania: Iguanidae). Their location at scale margins and the direction of their bristles towards adjacent scales indicates that the cutaneous sensory organs play an important role in proprioception during toepad deployment in Anolis.

The functional morphology of squamate fibrillar adhesive systems has been extensively investigated and has indirectly and directly influenced the design of synthetic counterparts. Not surprisingly, the structure and geometry of exemplar fibrils (setae) have been the subject of the bulk of the attention in such research, although variation in setal morphology along the length of subdigital adhesive pads has been implicated in the effective functioning of these systems. Adhesive setal field configuration has been described for several geckos, but that of the convergent Anolis lizards, comprised of morphologically simpler fibrils, remains largely unexplored. Here we examine setal morphology along the proximodistal axis of the digits of Anolis equestris and compare our findings to those for a model gecko, Gekko gecko. Consistent with previous work, we found that the setae of A. equestris are generally thinner, shorter, and present at higher densities than those of G. gecko and terminate in a single spatulate tip. Contrastingly, the setae of G. gecko are hierarchically branched in structure and carry hundreds of spatulate tips. Although the splitting of contacts into multiple smaller tips is predicted to increase the adhesive performance of a fiber compared to an unbranched one, we posited that the adhesive performance of G. gecko and A. equestris would be relatively similar when the configuration of the setal fields of each was accounted for. We found that, as in geckos, setal morphology of A. equestris follows a predictable pattern along the proximodistal axis of the pad, although there are several critical differences in the configuration of the setal fields of these two groups. Most notably, the pattern of variation in setal length of A. equestris is effectively opposite to that exhibited by G. gecko. This difference in clinal variation mirrors the difference in the direction in which the setal fields of anoles and geckos are peeled from the substrate, consistent with the hypothesis that biomechanical factors are the chief determinants of these patterns of variation. Future empirical work, however, is needed to validate this. Our findings introduce Anolis lizards as an additional source of inspiration for bio-inspired design and set the stage for comparative studies investigating the functional morphology of these convergent adhesive apparatuses. Such investigations will lead to an enhanced understanding of the interactions between form, function, and environment of fibril-based biological adhesive systems.

Ecologia trofca y morfologia de Anolis bartschi (Squamata: Dactyloidae) en el Parque Nacional Vinales, Cuba. Se conoce poco sobre la dieta de los anolinos cubanos. La morfologia en lagartos esta directamente relacionada con funciones ecologicas, como estrategias de forrajeo, competencias interespecifcas, o demanda energetica ligada a la reproduccion. Anolis bartschi es un endemico regional, restringido a las alturas carsicas del occidente de Cuba. Aqui ofrecemos datos nuevos sobre la ecologia trofca de esta especie y su relacion con la morfologia de su cabeza. Se capturaron 131 individuos adultos, donde los machos poseen mayor tamano del cuerpo y de la cabeza que las hembras. La mayoria de los individuos tenia presas en el estomago. Los machos consumiran mas presas que las hembras, pero de menor tamano que estas. Se detecto solapamiento trofco en epoca de seca, pero hubo segregamiento trofco en lluvia. Hymenoptera fue la presa mas consumida por ambos sexos. Ademas, las hembras consumieron mas Blattodea y Coleoptera, mientras los machos mas Diptera. Esta especie prefere presas sedentarias en lugar de moviles. Sugerimos que esta especie prefere presas sedentarias a presas moviles. De acuerdo a nuestros datos y las observaciones de campo, A. bartschi es un forrajeador bimodal, aunque se recomiendan futuras investigaciones sobre la variacion temporal (diaria y anual) en la dieta de esta especie para una apropiada clasifcacion sobre su actividad forrajeadora

Vertebrate predation by invertebrates has been classically underexplored and thus underestimated, despite the fact that many arthropods consume vertebrates. To shed some light on the relevance that spider predation may have upon lizards in the Neotropical and Andean regions, we compiled the available information in the literature on this trophic interaction. We found 50 reports of spiders consuming lizards in these regions, and the 88% of these were from the Neotropical region. Spiders belong to eight families, but Ctenidae and Theraphosidae were the most frequently reported predators. Lizards belong to 12 families, and the most commonly consumed species corresponded to the families Dactyloidae (all Anolis lizards), Gymnophthalmidae, and Sphaerodactylidae. Data suggest trophic spider–lizard associations between Ctenidae and Dactyloidae, followed by Theraphosidae and Liolaemidae. The body sizes of the spiders and lizards showed a positive relationship, and spiders were smaller than their prey. We conclude that various spider taxa can be considered lizard predators and they may be ecologically important in the Neotropical and Andean regions. However, spiders of prime predation relevance seem to be those of the Ctenidae and Theraphosidae families., In this study, we analyze for the first time the lizard consumption by spiders in the Neotropical and Andean regions, showing that the families Ctenidae and Theraphosidae may be relevant lizard predators in these areas. In addition, we provide the first report on spider predation upon a vertebrate taxon in the Andean region.

Native Green Anoles, Anolis carolinensis, and invasive Brown Anoles, Anolis sagrei, are commonly found in Florida and may be infected with the malarial parasite, Plasmodium floridense. Because no studies have directly addressed health effects of the parasite on Florida anoles, we collected blood smears of infected and uninfected anoles from Central and Southwest Florida and compared the overall leukocyte (WBC) counts, eosinophil counts, and heterophil/lymphocyte ratios. Eosinophils are generally elevated in response to protozoal infection and heterophil/lymphocyte ratios are often altered due to stress. A generalized linear model that tested contributions to erythrocyte/leukocyte ratios included infection status and locality as significant factors. We found significant differences in WBC counts between infected and uninfected lizards in Central Florida but not in Southwest Florida. Central Florida anoles also had higher mean WBC counts than Southwest Florida anoles. We did not detect significant differences in eosinophil counts or H/L ratios related to infection status. Our project is the first to examine leukocyte effects of Plasmodium infection in anoles and to provide leukocyte profiles of Anolis lizards. It appears that infected anoles sustain some negative immunological effects, at least in Central Florida. The differences in regions may be caused by the fact that Central Florida anoles still are under continuous interspecific competition whereas the Southwest Florida Brown Anoles are not because of low populations of Green Anoles. Additional studies that address leukocyte levels related to Plasmodium infection are needed to tease out the health and fitness effects on the lizards of Florida.

New localities and distribution models inform the conservation status of the endangered lizard Anolis guamuhaya (Squamata: Dactyloidae) from central Cuba. Anolis guamuhaya is known from seven localities restricted to the Guamuhaya Massif in central Cuba and is always associated with mountane ecosystems above 300 m a.s.l. Previous evaluations of the conservation status of the species based on the estimated number of mature individuals have categorized the anole as Endangered. Eight new records of A. guamuhaya are provided here. These double the number of known localities, and two represent the first records of the species in lowland areas, apart from the Guamuhaya Massif. The new records extend the elevational range of the species from 15 m to above 1000 m. We used ecological niche modeling based on all of the locality records, along with what we considered the most appropriate IUCN criteria according to the available information (Criterion B) to reevaluate the conservation status of the species. These new records of A. guamuhaya increase its area of occupancy up to a total of 60 km2 , and its extent of occurrence up to 648 km2 . Despite this increase in geographic range, the species meets the IUCN criteria in the category of Endangered. We used ecological niche modeling to predict possible trends for the species under differing scenarios of global climate change, all of which portend a drastic reduction in area climatically suitable for A. guamuhaya.

Cryptic species – genetically distinct species that are morphologically difficult to distinguish – present challenges to systematists. Operationally, cryptic species are very difficult to identify and sole use of genetic data or morphological data can fail to recognize evolutionarily isolated lineages. We use morphometric data to test species boundaries hypothesized with genetic data in the North Caribbean bark anole (Anolis distichus), a suspected species complex. We use univariate and multivariate analyses to test if candidate species based on genetic data can be accurately diagnosed. We also test alternative species delimitation scenarios with a model fitting approach that evaluates normal mixture models capable of identifying morphological clusters. Our analyses reject the hypothesis that the candidate species are diagnosable. Neither uni- nor multivariate morphometric data distinguish candidate species. The best-supported model included two morphological clusters; however, these clusters were uneven and did not align with a plausible species divergence scenario. After removing two related traits driving this result, only one cluster was supported. Despite substantial differentiation revealed by genetic data, we recover no new evidence to delimit species and refrain from taxonomic revision. This study highlights the importance of considering other types of data along with molecular data when delimiting species.

The composition of macro endoparasites associated with the lizard Norops fuscoauratus (Squamata) was analysed in two localities in the Atlantic Forest on the northeast of Brazil between December 2012 and July 2015. 74 specimens of N. fuscoauratus were examined and five species of helminths were reported, being: (a) for the population of Pernambuco: Cystacanth (Prevalence=37.5%), Physaloptera retusa Rudolphi, 1819 (Prevalence=4.16%), larva of flatworm (Prevalence=2.08%), Rhabdias sp. (Prevalence=2.08%) and Strongyluris oscari Travassos, 1923 (Prevalence=2.08%), and (b) of Alagoas: S. oscari (Prevalence=17.85%) and Rhabdias sp. (Prevalence=3.57%). The differences in the composition of endoparasites in the two populations are attributed to individualities of environments occupied by the lizards. The collection period does not influence the abundance of parasites, but when associated with sex, there was a positive correlation with the abundance of helminths, with more females than males being infected with parasites in the rainy season. Resumo A composição de macro endoparasitas associada com o lagarto Norops fuscoauratus (Squamata) foi analisada em duas localidades da Mata Atlântica no nordeste do Brasil, entre dezembro de 2012 e julho de 2015. 74 espécimes foram examinados e cinco espécies de helmintos foram encontradas, sendo: (a) para a população de Pernambuco: Cistacanto (Prevalência=37.5%), Physaloptera retusa Rudolphi, 1819 (Prevalência=4.16%), larva de platelminto (Prevalência=2.08%), Rhabdias sp. (Prevalência =2.08%) e Strongyluris oscari Travassos, 1923 (Prevalência =2.08%) e (b) Alagoas: S. oscari (Prevalência=17.85%) e Rhabdias sp. (Prevalência =3.57%). As diferenças na composição dos endoparasitas nas duas populações pode ser atribuída as individualidades dos ambientes ocupados por esses lagartos. O período de coleta não influenciou na abundância de parasitas, mas quando associado com o sexo, houve uma correlação positiva com a abundância de helmintos, com mais fêmeas do que machos, infectadas na estação chuvosa.

Thermal ecology and microhabitat use by lizards are affected directly and indirectly by intrinsic (e.g., sex and body size) and extrinsic (e.g., environmental seasonality, vegetation cover, and wind speed) factors. Herein we evaluate the effect of seasonality and sex on field thermal ecology and microhabitat utilization of a population of Anolis carlliebi inhabiting a xeric scrubland in Central Mexico. Lizards were found primarily in full and filtered sun conditions, strongly associated, respectively, with inflorescences and leaves of Agave stricta. Males were slightly larger than females. Mean field body temperature did not differ between sexes within seasons, but males used higher perches than females in the warm season. We observed a seasonal shift in body temperature and perch use in males, which exhibited higher body temperatures and occupied wider and higher perches in the warm season. Female body temperature and perch use did not differ between seasons. This study reports the first case of a population of Anolis specializing in a species of Agave as microhabitat. These results add to our understanding of thermoregulation and microhabitat use of mainland species of Anolis since they face different environmental pressures than Caribbean species.

We revise the species of Audantia, a genus of dactyloid lizards endemic to Hispaniola. Based on our analyses of morphological and genetic data we recognize 14 species in this genus, four of which we describe as new species (A. aridius sp. nov., A. australis sp. nov., A. higuey sp. nov., and A. hispaniolae sp. nov.), and two are resurrected from the synonymy of A. cybotes (A. doris comb. nov., A. ravifaux comb. nov.). Also, we place Anolis citrinellus Cope, 1864 in the synonymy of Ctenonotus distichus (Cope, 1861); Anolis haetianus Garman, 1887 in the synonymy of Audantia cybotes (Cope, 1863); and Anolis whitemani Williams, 1963 in the synonymy of Audantia saxatilis (Mertens, 1938). Finally, we designate a lectotype for Anolis cybotes Cope, 1863, and for Anolis riisei Reinhardt & Lütken, 1863. Our main focus is on the populations of anoles formerly referred to as Audantia cybotes which we demonstrate to be a complex of seven distinct species. For these seven species we provide a standardized description of external morphology, color descriptions in life, color photographs in life, description and illustration of hemipenis morphology (if available), distribution maps based on the specimens examined, comments on the conservation status, and natural history notes. Finally, we provide a dichotomous key for the identification of the 14 species of the genus Audantia occuring on Hispaniola.

Juvenile growth rates are thought to be restricted by available food resources. In animals that grow throughout the year, such as tropical lizards, growth is therefore predicted to be faster during the rainy season. We test this prediction using a population of Anolis nebulosusby describing the growth trajectories of both sexes using nonlinear regression models, and we then correlate the growth rates of individuals with food available in the environment, precipitation, and temperature. The Von Bertalanffy model fits the growth rates of the females better, while the logistic-by-length model fits the males better. According to both models, the males grew faster than females, reaching slightly smaller sizes at adulthood. Males reached sexual maturity when 35 mm long, at an age of seven months, and females matured at 37 mm (SVL), taking nine months to reach this size. In 1989, juvenile males and females grew more in both seasons (rainy and dry) than adults; for 1990, there were no differences by season or between age classes. These results are interesting since in the 1989 and 1990 rainy seasons, practically the same orders of prey and the greatest abundance of prey available in the environment were registered. A possible explanation could be that predation was more intense in 1990 than in 1989. There is little evidence that food, temperature, and humidity affect growth rates of A. nebulosus, refuting our predictions. This is mainly due to the low variation in growth observed in 1990. Therefore we think that the growth of this species reflects a complex combination of ecological and genetic factors.

Sexual dimorphism in body size, shape and colour is widespread in lizards as well as in other taxa. In the context of sexual selection, these sex differences are commonly manifested as exaggerated ...

Pleurodont lizards are characterized by an ancient system of sex chromosomes. Along with stability of the central component of the system (homologous to the X chromosome of Anolis carolinensis [Dactyloidae], ACAX), in some genera the ancestral sex chromosomes are fused with microautosomes, forming neo-sex chromosomes. The genus Ctenonotus (Dactyloidae) is characterized by multiple X1X1X2X2/X1X2Y sex chromosomes. According to cytogenetic data, the large neo-Y chromosome is formed by fusion of the ancestral Y chromosome with 2 microautosomes (homologous to ACA10 or ACA11 and ACA12), the X1 chromosome is formed by fusion of the ancestral X chromosome with the autosome homologous to ACA10 or ACA11, and the X2 chromosome is homologous to autosome ACA12. To determine more precisely the content and evolution of the Ctenonotus sex chromosomes, we sequenced flow-sorted chromosomes (both sex chromosomes and microautosomes as control) of 2 species with a similar system: C. pogus and C. sabanus. Our results indicate that the translocated part of the X1 is homologous to ACA11, X2 is homologous to ACA12, and the Y contains segments homologous to both ACA11 and ACA12. Molecular divergence estimates suggest that the ancestral X-derived part has completely degenerated in the Y of Ctenonotus, similar to the degeneration of the Norops sagrei Y chromosome (Dactyloidae). The newly added regions show loss of DNA content, but without degeneration of the conserved regions. We hypothesize that the translocation of autosomal blocks onto sex chromosomes facilitated rapid degeneration of the pseudoautosomal region on the ancestral Y.

Meiotic recombination rates and patterns of crossover distributions along the chromosomes vary considerably even between closely related species. The adaptive significance of these differences is still unclear due to the paucity of empirical data. Most data on recombination come from mammalian species, while other vertebrate clades are poorly explored. Using immunolocalization of the protein of the lateral element of the synaptonemal complex (SYCP3) and the mismatch-repair protein MLH1, which marks mature recombination nodules, we analyzed recombination rates and crossover distribution in meiotic prophase chromosomes of the steppe agama (Trapelus sanguinolentus, Agamidae, Acrodonta, Iguania) and compared them with data obtained for the genus Anolis (Dactyloidae, Pleurodonta, Iguania). We found that, despite a smaller genome size, the total SC length and the MLH1 focus number per cell are much higher in the agama than in the anoles. The distributions of the MLH1 foci in the agama are multimodal in larger chromosomes and bimodal in smaller chromosomes without a significant centromere effect, resembling the patterns known for birds. A possible relationship between karyotype remodeling and the evolution of recombination in Iguania is discussed.

We describe a new species of Anolis lizard from the Pacific slopes of the Andes of southwestern Ecuador at elevations between 372–1,000 m. The new species belongs to the Dactyloa clade and may be distinguished from other Anolis by size, external anatomy, mitochondrial DNA divergence, and dewlap color. Based on phylogenetic analyses of mitochondrial and nuclear DNA sequence data, we found that the new species is sister to A. fraseri in a clade composed primarily of large Dactyloid species. The new species is known from a protected area in southern Ecuador, Buenaventura Reserve, which suggests that at least some its populations are well protected.

Anolis nemonteae sp. nov. (Figs 1, 2, 3, 4, 5, 6) Proposed standard English name: Star anoles Proposed standard Spanish name: Anolis de las estrellas Holotype. QCAZ 14595 (Figs 1, 2, 3), adult female, Ecuador, El Oro province, Buenaventura Reserve, 3.654 S, 79.777 W, WGS 84, 417 m, 30 January 2016, collected by Andrea Narváez, Sebastián Valverde, Keyko Cruz, and David Reyes. Paratypes (N=12, Figs 3, 4). ECUADOR: El Oro: QCAZ 14317 (adult female), 14318 (juvenile female), 14431 (juvenile female), 14432 (adult female), 13.2 km SW Piñas on highway, 3.662 S, 79.760 W, 754 m, 11 January 2016, collected by Fernando Ayala, Steven Poe, and Chris Anderson; QCAZ 14596 (adult male), same collection data as holotype; QCAZ 14597 (juvenile male), Buenaventura Reserve, 3.650 S, 79.780 W, 372 m; QCAZ 14660 (female hatchling), Buenaventura Reserve, 3.653 S, 79.766 W, 578 m, 30 January 2016; DHMECN 4132 (juvenile male), Buenaventura Reserve, 3.645 S, 79.763 W, 800 m, 7 February 2006, collected by Mery Juiña; DHMECN 7687 (adult male), same collection data as DHMECN 4132, 18 January 2010, collected by Marco Reyes-Puig, and Michael Harvey; DHMECN 11543 (juvenile male), Buenaventura Reserve, 3.667 S, 79.766 W, 1,000 m, 31 December 2014, collected by Juan Carlos Sánchez, Karem López, Luis Oyagata, and Paúl Guerrero; JMG 0484 (adult male), 0485 (adult female), Buenaventura Lodge, 3.653 S, 79.768 W, 520 m, 6 January 2017, collected by Paulina Romero. Diagnosis. The new species belongs to the Megaloa clade of the latifrons series of Dactyloa (Castañeda & de Queiroz 2013; Prates et al. 2020) based on the phylogenetic tree presented in this study. Anolis nemonteae sp. nov. differs from most species of the punctatus, heterodermus, and nasofrontalis series (Castañeda & de Queiroz 2013; Prates et al. 2020) in having relatively smaller head scales; from the roquet series (Castañeda & de Queiroz 2013) in possessing supraorbital semicircles separated from each other and the interparietal separated from the supraorbital semicircles; and from the aequatorialis series (Castañeda & de Queiroz 2013; Prates et al. 2020) in having wider toepads and larger dorsal head scales. The new species is most similar in external morphology to the other members of the latifrons series (A. agassizi Stejneger 1900, A. apollinaris Boulenger 1919, A. brooksi Barbour 1923, A. casildae Arosemena, Ibáñez & de Sousa 1991, A. danieli Williams 1988, A. fraseri, A. frenatus Cope 1899, A. ginaelisae Lotzkat, Hertz, Bienentreu & Köhler 2013, A. ibanezi Poe, Latella, Ryan & Schaad 2009b, A. insignis Cope 1871, A. kathydayae Poe & Ryan 2017, A. kunayalae, A. latifrons Berthold 1846, A. limon Velasco & Hurtado-Gómez 2014, A. maculigula Williams 1984a, A. maia Batista, Vesely, Mebert, Lotzkat & Köhler 2015, A. microtus Cope 1871, A. mirus Williams 1963, A. parilis, A. princeps Boulenger 1902, A. propinquus Williams 1984b, A. purpurescens Cope 1899, A. savagei Poe & Ryan 2017, and A. squamulatus Peters 1863). Anolis nemonteae can readily be distinguished from A. agassizi, A. apollinaris, A. casildae, A. frenatus, A. ginaelisae, A. ibanezi, A. latifrons, A. limon, A. maculigula, A. maia, A. princeps, A. purpurescens, and A. squamulatus by having shorter legs not reaching ear when adpressed against body (legs reaching to ear or beyond when adpressed against body); from A. purpurescens, A. ibanezi, A. maia and A. limon further by having a green-brown dorsal background (green); from A. danieli and A. propinquus by lacking elongated superciliaries (one elongated superciliary); from A. brooksi, A. insignis, A. microtus, A. savagei, and A. kathydayae, all from Costa Rica and Panama, by possessing weakly keeled ventral scales (smooth); from A. ginaelisae (Panama) by lacking enlarged postcloacal scales in males (present); from A. kunayalae, A. mirus and A. parilis by having a wide toe pad on fourth toe (narrow toe pad), subdigital pad under phalanx III projecting above the proximal end of phalanx IV (subdigital pad continuous or indistinct), 21–23 lamellae under phalanges II and III of fourth toe (11–15 lamellae), and distal phalanx including claw equal or smaller than phalanges II and III combined (longer distal phalanx; see Fig. 1 of Williams 1963). Anolis nemonteae sp. nov. is most similar morphologically to A. fraseri in having a large body size (SVL> 85 mm), a green-brown dorsal background, reddish brown iris, smooth head scales, weakly keeled ventral scales, and 21–23 (18–24 in A. fraseri) lamellae under phalanges II and III of fourth toe. The new species can be distinguished from A. fraseri (character states in parentheses) by having a creamish white dewlap skin with black blotches longitudinally arranged along yellow stripes in females (female dewlap orangish yellow anteriorly, without black blotches); bluish white dewlap skin with yellowish white scales and gold apicogorgetal scales in males (creamish white skin with yellow or greenish white scales; Fig. 5); dark brown dots on neck laterally and dorsally (neck dots absent, large dark blotches might be present; Figs 3 and 5); 7–11, mean = 9.29±1.25 SD scales between second canthals, (6–10, 7.72±1.02, z = 2.737, p = 0.006); 3–4, 3.29±0.49 scales between supraorbital semicircles (2–4, 2.68±0.57, z = -2.281, p = 0.023), 7–10, 8.86±1.07 supralabials counted to a point below center of eye (7–9, 7.69±0.55, z = 2.845, p = 0.004); 21–23, 21.71±0.95 lamellae under phalanges II–III of fourth toe (18–23, 20.32±1.09, z = -2.799, p = 0.005), snout length/SVL, 0.116–0.122, 0.119±0.002 (0.112–0.133, 0.123±0.005, t = 2.664, p = 0.013) interparietal length/SVL, 0.016–0.028, 0.023±0.004 (0.011–0.027, 0.018±0.004, t = -2.750, p = 0.011), humerus length/ SVL, 0.131–0.184, 0.159±0.016 (0.155–0.204, 0.171±0.011, t = 2.380, p = 0.025), foot length/SVL 0.259–0.282, 0.272±0.009 (0.262–0.310, 0.287±0.014, t = 2.627, p = 0.014), and fourth toe length/SVL 0.151–0.177, 0.162±0.009 (0.168–0.203, 0.181±0.010, t = 4.287, p Anolis nemonteae sp. nov. differs from A. fraseri in lacking enlarged postcloacals in males (present, sometimes inconspicuous), and genetic distances between these species range between 0.03–0.05 (ND2) and 0.05–0.06 (COI). In addition to the above diagnostic traits, the unique male and female dewlap color patterns of A. nemonteae sp. nov. distinguish it from all other Ecuadorian Anolis of the Megaloa clade (Fig. 5). Description of the holotype (scores for paratypes in parentheses). Snout to vent length 92.4 mm (88.4−115.2 mm); tail length 191.5 mm (200.6−263.1 mm); head length 23.9 mm (23.6−31.3 mm); head width 14.7 mm (14.8−16.9 mm); head height 11.9 mm (12.1−14.4 mm); snout length 11.3 mm (10.4−13.4 mm); interorbital length 4.8 (4.4−5.8 mm); interparietal length 2.3 mm (1.8−2.9 mm); ear height 2.1 mm (1.7−2.5 mm); femur length 22.6 mm (20.5−27.4 mm); tibia length 18.6 mm (17.1−22.7 mm); foot length 25.6 mm (24.2−29.8 mm); humerus length 17.0 mm (13.5−19.2 mm); ulna length 13.3 mm (11.6−16.0 mm); hand length 14.5 mm (13.7−16.5 mm); fourth toe length 16.4 mm (14.3−19.2 mm); fourth toe width 1.8 mm (1.9−2.6 mm); dewlap length 28.8 mm (29.5−49.9 mm); and dewlap height 9.8 mm (6.3−15.2 mm). Head scales in supraocular disc and frontal region smooth (smooth or rugose); eleven (7−10) scales between second canthals; seven (6−9) scales bordering the rostral posteriorly; circumnasal separated by one (1−2) scale from rostral; supraorbital semicircles separated by three (3−4) scales; supraocular disc with four (3–4) enlarged scales; supraocular edge continuous; superciliaries in two series: first series without rectangular superciliary anteriorly (sometimes present) followed by gradually smaller scales, and second series with squarish scales; five (6−7) loreal rows;>15 loreal scales; midsnout without parallel scale rows (absent or weakly present); rostral with smooth dorsal edge; frontal region of head with a depression; rostral even with mental (even or slight overlapping); interparietal larger (smaller or larger) than ear opening, separated by three (1−3) scales from semicircles; ear oval with normal edge; transparent scales in lower eyelids absent; preoccipital scale absent; suboculars in contact with supralabials; ten (7−9) supralabials counted up to a point below center of eye; six (5−7) postmentals; no enlarged sublabials in contact with infralabials; mental divided partially; mental extending farther back posteriorly than rostral along edge of mouth (even or rostral extending farther than mental or mental extending farther back posteriorly than rostral); and posterior edge of mental straight (straight or slight concave). Low nuchal crest formed by continuous series of small conical scales; low middorsal crest formed by continuous series of triangular scales (crest present in adults of both sexes); dorsal scales slightly keeled anteriorly and smooth posteriorly (smooth or keeled anteriorly and keeled or smooth posteriorly); two (absent or two) vertebral rows slightly larger than flank scales; eight (9−11) middorsal scales in a longitudinal segment representing 5% of SVL; flank scales smooth (smooth, rugose or weakly keeled), homogeneous in size, and barely separated by skin; ventral scales smaller than dorsals (ventrals larger than dorsals), slightly keeled, subimbricate, with round posterior edge (round or rectangular), and arranged in diagonal rows; ten (7−9) midventral scales in a longitudinal segment representing 5% of SVL; inconspicuous axillary pocket present (present or absent). Toepads slightly overlapping distal phalanx in all toes; twenty-one (22−23) lamellae under phalanges II and III of fourth toe; supradigitals with multiple keels; tail crest absent; tail round (round or laterally compressed), with a single row of middorsal scales; insolitus tail (Poe 2004) absent; enlarged postcloacal scales absent (in males and females); hindlimb reaching posterior to ear when adpressed against body. Nuchal and dorsal folds present; dewlap large, extending posteriorly behind forelimbs (in males and females), with five (4−6) longitudinal rows of four (3–5) elongate scales each, smaller than ventrals, and separated by naked skin. Inter- and intraspecific variation in morphological characters in Anolis nemonteae is presented in Tables 2 and 3, respectively. Color in life. Holotype, adult female QCAZ 14595 (Fig. 3A, B, C, undisturbed color pattern): dorsum of head, body, limbs and tail pale yellowish green; dorsum of body with three broad, dark brown transverse bands extending onto flanks; dorsal surfaces of limbs and tail with dark brown transverse bands; palpebral scales yellowish green; flanks of neck and body with dark brown spots; ventral aspect of head, body, limbs, and tail cream; iris pale reddish brown; throat lining black; tongue yellow; edge of mouth including jaw hinges white; dewlap skin cream with black blotches, mostly arranged more or less longitudinally along yellow stripes (Fig. 5B); scales of dewlap yellowish white. mean and standard deviation are given. When stressed, adult females QCAZ 14594 and JMG0485 (Fig. 3D) turned dorsal background of head, body, limbs and tail yellowish brown or dark brown, respectively. Juvenile female QCAZ 14431 (Fig. 3E, stressed specimen): general color pattern similar to holotype, but dewlap skin dirty white with broad yellow stripes and elongate black blotches; gorgetals, sternals and marginals yellowish white; ventral aspect of head, body, limbs, and tail grayish cream with brown dots. Hatchling female QCAZ 14660 (Fig. 3F, stressed specimen): general color pattern similar to female QCAZ 14594, but dorsum of head, body, limbs and tail pale brown; dorsum of body with four dark brown transverse bands extending onto flanks. Adult male QCAZ 14596 (Fig. 3G, H, I, undisturbed color pattern): dorsum of body, limbs, and tail brown; dorsum of head light grayish turquoise with orange and red spots; dorsum of body with four broad, dark brown transverse bands extending onto flanks, of which the two posteriormost merge ventrally; second anteriormost broad band bordered anteriorly by yellowish cream line; dorsum of limbs and tail with dark brown transverse bands; palpebral scales yellow; flanks of neck and body with dark brown spots; ventral aspect of head, body, limbs, and tail cream; iris reddish brown; throat lining black; tongue yellow; edge of mouth including jaw hinges white; dewlap skin solid bluish white with yellowish white scales and golden apicogorgetals (Fig. 5A). When stressed, adult male JMG 0484 (Fig. 3J, K) turned background of body, limbs, and tail light brown. Color in preservative. In preservative, the holotype (QCAZ 14595) has a light brown background (Figs 1, 2); otherwise, the color patterns in life (see above) and preservative are similar. Adult male QCAZ 14596 (Fig. 4 A, B, left side): dorsum of body, limbs, and tail brown; dorsum of body with four dark brown transverse broad bands extending onto flanks; dorsum of limbs and tail with dark brown transverse bands; flanks of neck and body with dark brown spots; ventral aspect of head, body, and limbs grayish cream; tail dark brown with cream base; dewlap skin solid white with cream scales. Adult males JMG 0484 and DHMECN 7687 (Fig. 4 A, B, center and right side): general color pattern similar to male QCAZ 14596, but vertebral region of dorsum with dark brown spots; dorsum of DHMECN 7687 with three brown transverse bands of which the second one is bordered anteriorly by a cream stripe, which is discontinuous medially, where each half projects anteriorly. Adult female JMG 0485 (Fig. 4 C, D, center): dorsum of head, body, limbs and tail brown; dorsum of body with three broad, dark brown transverse bands extending onto flanks; vertebral region of dorsum with dark brown spots; dorsum of limbs and tail with dark brown transverse bands; flanks of neck and body with dark brown spots; ventral aspect of head, body, limbs, and tail dirty cream; dewlap skin creamish white with black blotches; scales of dewlap creamish white. Adult female QCAZ 14432 (Fig. 4 C, D, right side) differs from JMG 0485 in having faint dorsal transverse bands and small dark brown spots on limbs ventrally. Female hatchling QCAZ 14660 (Fig. 4 C, D, left side) differs in having four dark brown transverse bands (discontinuous medially) on ventral aspect of head and dark brown transverse bands ventrally on limbs. Hemipenis. Partially everted hemipenis small (7.3 mm total length, 2.9 mm truncus length) and slightly bilobate; sulcus spermaticus unforked, bordered by well-developed lips and opening into a single, smooth apical area; lobes with small calyces on asulcate side; fleshy projection on lobular crotch (crotch flap) on asulcate surface; truncus with transverse and lateral folds (Fig. 6). Distribution and natural history. Anolis nemonteae occurs on the Pacific slopes of the Andes in southern Ecuador, El Oro province, between 372−1,000 m (Fig. 7). This species occurs in the Evergreen Foothills Montane Forest of Catamayo-Alamor (Ministerio del Ambiente del Ecuador 2013). Specimens of Anolis nemonteae were collected mainly in secondary forest and along the edge of roads (Fig. 8). All individuals were found at night between 20:00 and 01:00 h sleeping horizontally on branches of trees, or leaves of bananas or Panama hat plants, 2.5–6.9 m above ground. A male and a female (JMG 0484, 0485) were found sleeping on banana leaves 10 m away from each other, 2.5 m above the ground. Anolis nemonteae occurs in sympatry with A. binotatus Peters 1863 and A. fasciatus at the type locality (Yánez-Muñoz et al. 2013, Garzón et al. 2019). A captive female (QCAZ 14594 [tissue sample], SVL = 93.21 mm) that was subsequently released laid one egg (22.3 x 14.4 mm) on 30 January 2016. The egg was incubated in perlite at 19°C and 85% of relative humidity. After an incubation period of 129 days, a female (Fig. 3F; QCAZ 14660, SVL = 32.9 mm, weight = 2.3 g) hatched. Conservation. The known distribution area of Anolis nemonteae has suffered from dramatic deforestation (Tapia-Armijos et al. 2015). However, most individuals of A. nemonteae were collected within the Buenaventura Reserve, which suggests that at least some of its populations are well protected. Because of the small known distribution (Fig. 7) and lack of additional data, we suggest assigning A. nemonteae to the Data Deficient category according to IUCN (2012) guidelines. Etymology. The specific epithet nemonteae is a noun in the genitive case and is a patronym for Nemonte Nenquimo, indigenous activist who led a successful campaign and legal action that protected 500,000 acres of Amazonian rainforest and Waorani territory from oil extraction in Ecuador. Nemonte means ‘many stars’ in Wao Tereo language. Nemonte Nenquimo’s work has been recognized worldwide. In 2020, she was awarded the prestigious Goldman Prize and was listed among the 100 most influential people of the year by the Time Magazine. Here we honor Nemonte Nenquimo for her braveness and determination to protect natural forests and their inhabitants. Phylogenetic relationships. Data partitions and models of evolution are presented in Table 4. Both ML and Bayesian analyses positioned Anolis nemonteae sp. nov. within a clade highly congruent with the Megaloa clade of Castañeda & de Queiroz (2013) with maximum support (Fig. 9). The difference to the Megaloa clade as originally defined is that Anolis ibanezi, A. kunayalae, and A. parilis were also included (Fig. 9). As expected, A. ginaelisae and A. maia, both described more recently, were also included in this clade (Lotzkat et al. 2013, Batista et al. 2015). Both ML and Bayesian tree topologies for the Megaloa clade are identical, except for the sister species of A. maculigula (A. casildae and A. apollinaris, respectively). The sister species of A. nemonteae sp. nov. is A. fraseri (PP = 0.98, BS = 65), and together they form a clade sister to A. parilis (PP = 1, BS = 84). Interspecific genetic distances among sampled species of the Megaloa clade range from 0.02 (A. fraseri / A. parilis, A. latifrons / A. princeps) to 0.23 (A. latifrons /

Anolis nemonteae sp. nov. (Figs 1, 2, 3, 4, 5, 6) Proposed standard English name: Star anoles Proposed standard Spanish name: Anolis de las estrellas Holotype. QCAZ 14595 (Figs 1, 2, 3), adult female, Ecuador, El Oro province, Buenaventura Reserve, 3.654 S, 79.777 W, WGS 84, 417 m, 30 January 2016, collected by Andrea Narváez, Sebastián Valverde, Keyko Cruz, and David Reyes. Paratypes (N=12, Figs 3, 4). ECUADOR: El Oro: QCAZ 14317 (adult female), 14318 (juvenile female), 14431 (juvenile female), 14432 (adult female), 13.2 km SW Piñas on highway, 3.662 S, 79.760 W, 754 m, 11 January 2016, collected by Fernando Ayala, Steven Poe, and Chris Anderson; QCAZ 14596 (adult male), same collection data as holotype; QCAZ 14597 (juvenile male), Buenaventura Reserve, 3.650 S, 79.780 W, 372 m; QCAZ 14660 (female hatchling), Buenaventura Reserve, 3.653 S, 79.766 W, 578 m, 30 January 2016; DHMECN 4132 (juvenile male), Buenaventura Reserve, 3.645 S, 79.763 W, 800 m, 7 February 2006, collected by Mery Juiña; DHMECN 7687 (adult male), same collection data as DHMECN 4132, 18 January 2010, collected by Marco Reyes-Puig, and Michael Harvey; DHMECN 11543 (juvenile male), Buenaventura Reserve, 3.667 S, 79.766 W, 1,000 m, 31 December 2014, collected by Juan Carlos Sánchez, Karem López, Luis Oyagata, and Paúl Guerrero; JMG 0484 (adult male), 0485 (adult female), Buenaventura Lodge, 3.653 S, 79.768 W, 520 m, 6 January 2017, collected by Paulina Romero. Diagnosis. The new species belongs to the Megaloa clade of the latifrons series of Dactyloa (Castañeda & de Queiroz 2013; Prates et al. 2020) based on the phylogenetic tree presented in this study. Anolis nemonteae sp. nov. differs from most species of the punctatus, heterodermus, and nasofrontalis series (Castañeda & de Queiroz 2013; Prates et al. 2020) in having relatively smaller head scales; from the roquet series (Castañeda & de Queiroz 2013) in possessing supraorbital semicircles separated from each other and the interparietal separated from the supraorbital semicircles; and from the aequatorialis series (Castañeda & de Queiroz 2013; Prates et al. 2020) in having wider toepads and larger dorsal head scales. The new species is most similar in external morphology to the other members of the latifrons series (A. agassizi Stejneger 1900, A. apollinaris Boulenger 1919, A. brooksi Barbour 1923, A. casildae Arosemena, Ibáñez & de Sousa 1991, A. danieli Williams 1988, A. fraseri, A. frenatus Cope 1899, A. ginaelisae Lotzkat, Hertz, Bienentreu & Köhler 2013, A. ibanezi Poe, Latella, Ryan & Schaad 2009b, A. insignis Cope 1871, A. kathydayae Poe & Ryan 2017, A. kunayalae, A. latifrons Berthold 1846, A. limon Velasco & Hurtado-Gómez 2014, A. maculigula Williams 1984a, A. maia Batista, Vesely, Mebert, Lotzkat & Köhler 2015, A. microtus Cope 1871, A. mirus Williams 1963, A. parilis, A. princeps Boulenger 1902, A. propinquus Williams 1984b, A. purpurescens Cope 1899, A. savagei Poe & Ryan 2017, and A. squamulatus Peters 1863). Anolis nemonteae can readily be distinguished from A. agassizi, A. apollinaris, A. casildae, A. frenatus, A. ginaelisae, A. ibanezi, A. latifrons, A. limon, A. maculigula, A. maia, A. princeps, A. purpurescens, and A. squamulatus by having shorter legs not reaching ear when adpressed against body (legs reaching to ear or beyond when adpressed against body); from A. purpurescens, A. ibanezi, A. maia and A. limon further by having a green-brown dorsal background (green); from A. danieli and A. propinquus by lacking elongated superciliaries (one elongated superciliary); from A. brooksi, A. insignis, A. microtus, A. savagei, and A. kathydayae, all from Costa Rica and Panama, by possessing weakly keeled ventral scales (smooth); from A. ginaelisae (Panama) by lacking enlarged postcloacal scales in males (present); from A. kunayalae, A. mirus and A. parilis by having a wide toe pad on fourth toe (narrow toe pad), subdigital pad under phalanx III projecting above the proximal end of phalanx IV (subdigital pad continuous or indistinct), 21–23 lamellae under phalanges II and III of fourth toe (11–15 lamellae), and distal phalanx including claw equal or smaller than phalanges II and III combined (longer distal phalanx; see Fig. 1 of Williams 1963). Anolis nemonteae sp. nov. is most similar morphologically to A. fraseri in having a large body size (SVL> 85 mm), a green-brown dorsal background, reddish brown iris, smooth head scales, weakly keeled ventral scales, and 21–23 (18–24 in A. fraseri) lamellae under phalanges II and III of fourth toe. The new species can be distinguished from A. fraseri (character states in parentheses) by having a creamish white dewlap skin with black blotches longitudinally arranged along yellow stripes in females (female dewlap orangish yellow anteriorly, without black blotches); bluish white dewlap skin with yellowish white scales and gold apicogorgetal scales in males (creamish white skin with yellow or greenish white scales; Fig. 5); dark brown dots on neck laterally and dorsally (neck dots absent, large dark blotches might be present; Figs 3 and 5); 7–11, mean = 9.29±1.25 SD scales between second canthals, (6–10, 7.72±1.02, z = 2.737, p = 0.006); 3–4, 3.29±0.49 scales between supraorbital semicircles (2–4, 2.68±0.57, z = -2.281, p = 0.023), 7–10, 8.86±1.07 supralabials counted to a point below center of eye (7–9, 7.69±0.55, z = 2.845, p = 0.004); 21–23, 21.71±0.95 lamellae under phalanges II–III of fourth toe (18–23, 20.32±1.09, z = -2.799, p = 0.005), snout length/SVL, 0.116–0.122, 0.119±0.002 (0.112–0.133, 0.123±0.005, t = 2.664, p = 0.013) interparietal length/SVL, 0.016–0.028, 0.023±0.004 (0.011–0.027, 0.018±0.004, t = -2.750, p = 0.011), humerus length/ SVL, 0.131–0.184, 0.159±0.016 (0.155–0.204, 0.171±0.011, t = 2.380, p = 0.025), foot length/SVL 0.259–0.282, 0.272±0.009 (0.262–0.310, 0.287±0.014, t = 2.627, p = 0.014), and fourth toe length/SVL 0.151–0.177, 0.162±0.009 (0.168–0.203, 0.181±0.010, t = 4.287, p Anolis nemonteae sp. nov. differs from A. fraseri in lacking enlarged postcloacals in males (present, sometimes inconspicuous), and genetic distances between these species range between 0.03–0.05 (ND2) and 0.05–0.06 (COI). In addition to the above diagnostic traits, the unique male and female dewlap color patterns of A. nemonteae sp. nov. distinguish it from all other Ecuadorian Anolis of the Megaloa clade (Fig. 5). Description of the holotype (scores for paratypes in parentheses). Snout to vent length 92.4 mm (88.4−115.2 mm); tail length 191.5 mm (200.6−263.1 mm); head length 23.9 mm (23.6−31.3 mm); head width 14.7 mm (14.8−16.9 mm); head height 11.9 mm (12.1−14.4 mm); snout length 11.3 mm (10.4−13.4 mm); interorbital length 4.8 (4.4−5.8 mm); interparietal length 2.3 mm (1.8−2.9 mm); ear height 2.1 mm (1.7−2.5 mm); femur length 22.6 mm (20.5−27.4 mm); tibia length 18.6 mm (17.1−22.7 mm); foot length 25.6 mm (24.2−29.8 mm); humerus length 17.0 mm (13.5−19.2 mm); ulna length 13.3 mm (11.6−16.0 mm); hand length 14.5 mm (13.7−16.5 mm); fourth toe length 16.4 mm (14.3−19.2 mm); fourth toe width 1.8 mm (1.9−2.6 mm); dewlap length 28.8 mm (29.5−49.9 mm); and dewlap height 9.8 mm (6.3−15.2 mm). Head scales in supraocular disc and frontal region smooth (smooth or rugose); eleven (7−10) scales between second canthals; seven (6−9) scales bordering the rostral posteriorly; circumnasal separated by one (1−2) scale from rostral; supraorbital semicircles separated by three (3−4) scales; supraocular disc with four (3–4) enlarged scales; supraocular edge continuous; superciliaries in two series: first series without rectangular superciliary anteriorly (sometimes present) followed by gradually smaller scales, and second series with squarish scales; five (6−7) loreal rows;>15 loreal scales; midsnout without parallel scale rows (absent or weakly present); rostral with smooth dorsal edge; frontal region of head with a depression; rostral even with mental (even or slight overlapping); interparietal larger (smaller or larger) than ear opening, separated by three (1−3) scales from semicircles; ear oval with normal edge; transparent scales in lower eyelids absent; preoccipital scale absent; suboculars in contact with supralabials; ten (7−9) supralabials counted up to a point below center of eye; six (5−7) postmentals; no enlarged sublabials in contact with infralabials; mental divided partially; mental extending farther back posteriorly than rostral along edge of mouth (even or rostral extending farther than mental or mental extending farther back posteriorly than rostral); and posterior edge of mental straight (straight or slight concave). Low nuchal crest formed by continuous series of small conical scales; low middorsal crest formed by continuous series of triangular scales (crest present in adults of both sexes); dorsal scales slightly keeled anteriorly and smooth posteriorly (smooth or keeled anteriorly and keeled or smooth posteriorly); two (absent or two) vertebral rows slightly larger than flank scales; eight (9−11) middorsal scales in a longitudinal segment representing 5% of SVL; flank scales smooth (smooth, rugose or weakly keeled), homogeneous in size, and barely separated by skin; ventral scales smaller than dorsals (ventrals larger than dorsals), slightly keeled, subimbricate, with round posterior edge (round or rectangular), and arranged in diagonal rows; ten (7−9) midventral scales in a longitudinal segment representing 5% of SVL; inconspicuous axillary pocket present (present or absent). Toepads slightly overlapping distal phalanx in all toes; twenty-one (22−23) lamellae under phalanges II and III of fourth toe; supradigitals with multiple keels; tail crest absent; tail round (round or laterally compressed), with a single row of middorsal scales; insolitus tail (Poe 2004) absent; enlarged postcloacal scales absent (in males and females); hindlimb reaching posterior to ear when adpressed against body. Nuchal and dorsal folds present; dewlap large, extending posteriorly behind forelimbs (in males and females), with five (4−6) longitudinal rows of four (3–5) elongate scales each, smaller than ventrals, and separated by naked skin. Inter- and intraspecific variation in morphological characters in Anolis nemonteae is presented in Tables 2 and 3, respectively. Color in life. Holotype, adult female QCAZ 14595 (Fig. 3A, B, C, undisturbed color pattern): dorsum of head, body, limbs and tail pale yellowish green; dorsum of body with three broad, dark brown transverse bands extending onto flanks; dorsal surfaces of limbs and tail with dark brown transverse bands; palpebral scales yellowish green; flanks of neck and body with dark brown spots; ventral aspect of head, body, limbs, and tail cream; iris pale reddish brown; throat lining black; tongue yellow; edge of mouth including jaw hinges white; dewlap skin cream with black blotches, mostly arranged more or less longitudinally along yellow stripes (Fig. 5B); scales of dewlap yellowish white. mean and standard deviation are given. When stressed, adult females QCAZ 14594 and JMG0485 (Fig. 3D) turned dorsal background of head, body, limbs and tail yellowish brown or dark brown, respectively. Juvenile female QCAZ 14431 (Fig. 3E, stressed specimen): general color pattern similar to holotype, but dewlap skin dirty white with broad yellow stripes and elongate black blotches; gorgetals, sternals and marginals yellowish white; ventral aspect of head, body, limbs, and tail grayish cream with brown dots. Hatchling female QCAZ 14660 (Fig. 3F, stressed specimen): general color pattern similar to female QCAZ 14594, but dorsum of head, body, limbs and tail pale brown; dorsum of body with four dark brown transverse bands extending onto flanks. Adult male QCAZ 14596 (Fig. 3G, H, I, undisturbed color pattern): dorsum of body, limbs, and tail brown; dorsum of head light grayish turquoise with orange and red spots; dorsum of body with four broad, dark brown transverse bands extending onto flanks, of which the two posteriormost merge ventrally; second anteriormost broad band bordered anteriorly by yellowish cream line; dorsum of limbs and tail with dark brown transverse bands; palpebral scales yellow; flanks of neck and body with dark brown spots; ventral aspect of head, body, limbs, and tail cream; iris reddish brown; throat lining black; tongue yellow; edge of mouth including jaw hinges white; dewlap skin solid bluish white with yellowish white scales and golden apicogorgetals (Fig. 5A). When stressed, adult male JMG 0484 (Fig. 3J, K) turned background of body, limbs, and tail light brown. Color in preservative. In preservative, the holotype (QCAZ 14595) has a light brown background (Figs 1, 2); otherwise, the color patterns in life (see above) and preservative are similar. Adult male QCAZ 14596 (Fig. 4 A, B, left side): dorsum of body, limbs, and tail brown; dorsum of body with four dark brown transverse broad bands extending onto flanks; dorsum of limbs and tail with dark brown transverse bands; flanks of neck and body with dark brown spots; ventral aspect of head, body, and limbs grayish cream; tail dark brown with cream base; dewlap skin solid white with cream scales. Adult males JMG 0484 and DHMECN 7687 (Fig. 4 A, B, center and right side): general color pattern similar to male QCAZ 14596, but vertebral region of dorsum with dark brown spots; dorsum of DHMECN 7687 with three brown transverse bands of which the second one is bordered anteriorly by a cream stripe, which is discontinuous medially, where each half projects anteriorly. Adult female JMG 0485 (Fig. 4 C, D, center): dorsum of head, body, limbs and tail brown; dorsum of body with three broad, dark brown transverse bands extending onto flanks; vertebral region of dorsum with dark brown spots; dorsum of limbs and tail with dark brown transverse bands; flanks of neck and body with dark brown spots; ventral aspect of head, body, limbs, and tail dirty cream; dewlap skin creamish white with black blotches; scales of dewlap creamish white. Adult female QCAZ 14432 (Fig. 4 C, D, right side) differs from JMG 0485 in having faint dorsal transverse bands and small dark brown spots on limbs ventrally. Female hatchling QCAZ 14660 (Fig. 4 C, D, left side) differs in having four dark brown transverse bands (discontinuous medially) on ventral aspect of head and dark brown transverse bands ventrally on limbs. Hemipenis. Partially everted hemipenis small (7.3 mm total length, 2.9 mm truncus length) and slightly bilobate; sulcus spermaticus unforked, bordered by well-developed lips and opening into a single, smooth apical area; lobes with small calyces on asulcate side; fleshy projection on lobular crotch (crotch flap) on asulcate surface; truncus with transverse and lateral folds (Fig. 6). Distribution and natural history. Anolis nemonteae occurs on the Pacific slopes of the Andes in southern Ecuador, El Oro province, between 372−1,000 m (Fig. 7). This species occurs in the Evergreen Foothills Montane Forest of Catamayo-Alamor (Ministerio del Ambiente del Ecuador 2013). Specimens of Anolis nemonteae were collected mainly in secondary forest and along the edge of roads (Fig. 8). All individuals were found at night between 20:00 and 01:00 h sleeping horizontally on branches of trees, or leaves of bananas or Panama hat plants, 2.5–6.9 m above ground. A male and a female (JMG 0484, 0485) were found sleeping on banana leaves 10 m away from each other, 2.5 m above the ground. Anolis nemonteae occurs in sympatry with A. binotatus Peters 1863 and A. fasciatus at the type locality (Yánez-Muñoz et al. 2013, Garzón et al. 2019). A captive female (QCAZ 14594 [tissue sample], SVL = 93.21 mm) that was subsequently released laid one egg (22.3 x 14.4 mm) on 30 January 2016. The egg was incubated in perlite at 19°C and 85% of relative humidity. After an incubation period of 129 days, a female (Fig. 3F; QCAZ 14660, SVL = 32.9 mm, weight = 2.3 g) hatched. Conservation. The known distribution area of Anolis nemonteae has suffered from dramatic deforestation (Tapia-Armijos et al. 2015). However, most individuals of A. nemonteae were collected within the Buenaventura Reserve, which suggests that at least some of its populations are well protected. Because of the small known distribution (Fig. 7) and lack of additional data, we suggest assigning A. nemonteae to the Data Deficient category according to IUCN (2012) guidelines. Etymology. The specific epithet nemonteae is a noun in the genitive case and is a patronym for Nemonte Nenquimo, indigenous activist who led a successful campaign and legal action that protected 500,000 acres of Amazonian rainforest and Waorani territory from oil extraction in Ecuador. Nemonte means ‘many stars’ in Wao Tereo language. Nemonte Nenquimo’s work has been recognized worldwide. In 2020, she was awarded the prestigious Goldman Prize and was listed among the 100 most influential people of the year by the Time Magazine. Here we honor Nemonte Nenquimo for her braveness and determination to protect natural forests and their inhabitants. Phylogenetic relationships. Data partitions and models of evolution are presented in Table 4. Both ML and Bayesian analyses positioned Anolis nemonteae sp. nov. within a clade highly congruent with the Megaloa clade of Castañeda & de Queiroz (2013) with maximum support (Fig. 9). The difference to the Megaloa clade as originally defined is that Anolis ibanezi, A. kunayalae, and A. parilis were also included (Fig. 9). As expected, A. ginaelisae and A. maia, both described more recently, were also included in this clade (Lotzkat et al. 2013, Batista et al. 2015). Both ML and Bayesian tree topologies for the Megaloa clade are identical, except for the sister species of A. maculigula (A. casildae and A. apollinaris, respectively). The sister species of A. nemonteae sp. nov. is A. fraseri (PP = 0.98, BS = 65), and together they form a clade sister to A. parilis (PP = 1, BS = 84). Interspecific genetic distances among sampled species of the Megaloa clade range from 0.02 (A. fraseri / A. parilis, A. latifrons / A. princeps) to 0.23 (A. latifrons /, Published as part of Ayala-Varela, Fernando, Valverde, Sebastián, Poe, Steven, Narváez, Andrea E., Yánez-Muñoz, Mario H. & Torres-Carvajal, Omar, 2021, A new giant anole (Squamata: Iguanidae: Dactyloinae) from southwestern Ecuador, pp. 295-317 in Zootaxa 4991 (2) on pages 298-311, DOI: 10.11646/zootaxa.4991.2.4, http://zenodo.org/record/5092205, {"references":["Castaneda, M. R. & de Queiroz, K. (2013) Phylogeny of the Dactyloa clade of Anolis lizards: new insights from combining morphological and molecular data. Bulletin of the Museum of Comparative Zoology, 160, 345 - 398. https: // doi. org / 10.3099 / 0027 - 4100 - 160.7.345","Prates, I., Melo-Sampaio, P., de Queiroz, K., Carnaval, A., Rodrigues, M. & Drummond, L. (2020) Discovery of a new species of Anolis lizards from Brazil and its implications for the historical biogeography of montane Atlantic Forest endemics. Amphibia-Reptilia, 41 (1), 87 - 103. https: // doi. org / 10.1163 / 15685381 - 20191179","Stejneger, L. (1900) Descriptions of two new lizards of the genus Anolis from Cocos and Malpelo Islands. Bulletin of the Museum of Comparative Zoology at Harvard College, 36, 161 - 163.","Boulenger, G. A. (1919) Descriptions of two new lizards and a new frog from the Andes of Colombia. Procedings of the Zoological Society of London, 1919, 79 - 81. https: // doi. org / 10.1111 / j. 1096 - 3642.1919. tb 02114. x","Barbour, T. (1923) Notes on reptiles and amphibians from Panam. Occasional Papers of the Museum of Zoology, University of Michigan, 129, 1 - 16.","Arosemena, F. A., Ibanez, D. R. & De Sousa, F. (1991) Una nueva especie de Anolis (Squamata: Iguanidae) del grupo latifrons de Fortuna, Panam. Revista de Biologia Tropical, 39 (2), 255 - 262.","Williams, E. E. (1988). New or problematic Anolis from Colombia. V. Anolis danieli, a new species of the latifrons species group and a reassessment of Anolis apollinaris Boulenger, 1919. Breviora, 489, 1 - 25.","Cope, E. D. (1899) Contributions to the herpetology of New Granada and Argentina, with descriptions of new forms. Philadelphia Museum of Science Bulletin, 1, 1 - 19. https: // doi. org / 10.5962 / bhl. title. 54674","Lotzkat, S., Hertz, A., Bienentreu, J. - F. & Kohler, G. (2013) Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus. Zootaxa, 3626 (1), 1 - 54. https: // doi. org / 10.11646 / zootaxa. 3626.1.1","Poe, S., Latella, I. M., Ryan, M. J. & Schaad, E. W. (2009 b) A new species of Anolis lizard (Squamata, Iguania) from Panama. Phyllomedusa, 8 (2), 81 - 87. https: // doi. org / 10.11606 / issn. 2316 - 9079. v 8 i 2 p 81 - 87","Cope, E. D. (1871) Ninth contribution to the herpetology of tropical America. Proceedings of the Academy of Natural Science of Philadelphia, 23, 200 - 224.","Poe, S. & Ryan, M. J. (2017) Description of two new species similar to Anolis insignis (Squamata: Iguanidae) and resurrection of Anolis (Diaphoranolis) brooksi. Amphibian & Reptile Conservation, 11 (2), 1 - 16.","Berthold, A. A. (1846) Uber verschiedene neue oder seltene Reptilien aus Neu-Granada und Crustaceen aus China. Abhandlungen der Koniglichen Gesellschaft der Wissenschaften zu Gottingen 3, 3 - 32. https: // doi. org / 10.5962 / bhl. title. 5485","Velasco, J. A. & Hurtado-Gomez, J. P. (2014) A new green anole lizard of the Dactyloa clade (Squamata: Dactyloidae) from the Magdalena river valley of Colombia. Zootaxa, 3785 (2), 201 - 216. https: // doi. org / 10.11646 / zootaxa. 3785.2.4","Williams, E. E. (1984 a) New or problematic Anolis from Colombia. III. Two new semiaquatic anoles from Antioquia and Choco, Colombia. Breviora, 478, 1 - 22.","Williams, E. E. (1963) Studies on South American anoles. Description of Anolis mirus, new species from Rio San Juan, Colombia, with comment on digital dilation and dewlap as generic and specific characters in the anoles. Bulletin of the Museum of Comparative Zoology at Harvard, 129, 463 - 480.","Boulenger, G. A. (1902) Descriptions of new batrachians and reptiles from northwestern Ecuador. Annals and Magazine of Natural History, Series 7, 9 (49), 51 - 57. https: // doi. org / 10.1080 / 00222930208678538","Williams, E. E. (1984 b) New or problematic Anolis from Colombia. II. Anolis propinquus, another new species from the cloud forest of western Colombia. Breviora, 477, 1 - 7.","Peters, W. C. H. (1863) Uber einige neue Arten der Saurier-Gattung Anolis. Monatsberichte der Koniglichen Preussischen Akademie des Wissenschaften zu Berlin, 1863, 135 - 149.","Poe, S. (2004) Phylogeny of Anoles. Herpetological Monographs, 18, 37 - 89. https: // doi. org / 10.1655 / 0733 - 1347 (2004) 018 [0037: POA] 2.0. CO; 2","Yanez-Munoz, M. H., Morales, M., Reyes-Puig, M. & Meza-Ramos, P. A. (2013) Reserva Biologica Buenaventura: entre la transicion humeda tropical y la influencia tumbesina. In: MECN, Jocotoco & Ecominga (Eds.), Herpetofauna en areas prioriotarias para la conservacion: El sistema de Reservas Jocotoco y Ecominga. Serie de Publicaciones del Museo Ecuatoriano de Ciencias Naturales (MECN), Fundacion para la Conservacion Jocotoco, Fundacion Ecominga, Quito-Ecuador, Monografia 6, pp. 62 - 76.","Tapia-Armijos, M. F., Homeier, J., Espinosa, C. I., Leuschner, C. & de la Cruz, M. (2015) Deforestation and forest fragmentation in south Ecuador since the 1970 s - Losing a hotspot of biodiversity. PLoS ONE, 10 (11), e 0142359. https: // doi. org / 10.1371 / journal. pone. 0133701","IUCN (2012) IUCN Red List categories and criteria. Version 3.1. 2 nd Edition. IUCN Species Survival Commission, Gland, Switzerland and Cambridge, vi + 34 pp.","Batista, A., Vesely, M., Mebert. K., Lotzkat, S. & K ohler, G. (2015) A new species of Dactyloa from eastern Panama, with comments on other Dactyloa species present in the region. Zootaxa, 4039 (1), 57 - 84. https: // doi. org / 10.11646 / zootaxa. 4039.1.2","Fleishman, L. J, Leal, M. & Persons, M. H. (2009) Habitat light and dewlap color diversity in four species of Puerto Rican anoline lizards. Journal of Comparative Physiology A, 195, 1043 - 60. https: // doi. org / 10.1007 / s 00359 - 009 - 0478 - 8","Torres-Carvajal, O., Ayala-Varela, F., Lobos, S., Poe, S. & Narvaez, A. (2018) Two new Andean species of Anolis lizard (Iguanidae: Dactyloinae) from southern Ecuador. Journal of Natural History, 52, 1067 - 1089. https: // doi. org / 10.1080 / 00222933.2017.1391343"]}

Bergmann's rule predicts an inverse relationship between body size and environmental temperature in endotherms. The larger the organism, the lower the rate of heat loss by radiation; thus, larger organisms should be more likely to survive in colder places. This relationship is less clear in ectotherms. We tested Bergmann's hypothesis for a sample of 260 individuals of Anolis nebulosus from four Mexican populations ranging from a continental island to a forest at 2,100 m in elevation. Using analysis of variance, we tested for differences in body size (first principal component extracted from seven morphometric variables) among populations. Average body size was expected to be significantly larger in higher localities. Results did not conform to expectation; the island population was significantly larger in body size than the rest likely because of lower predation pressure. Also, we examined the relationship between body size, perch temperature (PT), and the difference between body temperature and PT. There were significant differences among populations in the range of PTs used. Individuals from highlands, where temperatures are colder and vary greatly, used a broader range of PTs and used perches that were colder than their body temperature. Only the largest individuals, mostly males, were able to use the coldest perches. In contrast, at lower sites, which had less thermal variation, lizards showed less variance in PTs, and there was no relationship between PT and size. In general, variation in PT and body size was smaller for females than for males. Results suggest a combination of heliothermy and heat conservation by size as regulatory strategies, and a trade-off between mating strategies and heat loss. For these populations, body size is one of a number of factors involved in temperature regulation; the particular combination of these factors on a population is probably a response to local environmental conditions.

A combined approach based on karyology and DNA taxonomy allowed us to characterize the taxonomic peculiarities in 10 Mesoamerican lizard species, belonging to six genera and five families, inhabiting two Biosphere Reserve in Chiapas, Mexico: La Sepultura Biosphere Reserve, and Montes Azules Biosphere. The karyotypes of four species, Phyllodactylus sp. 3 ( P. tuberculosus species group) (2n = 38), Holcosus festivus (Lichtenstein et von Martens, 1856) (2n = 50), Anolis lemurinus Cope, 1861 (2n = 40), and A. uniformis Cope, 1885 (2n = 29–30) are described for the first time, the last one showing a particular X1X1X2X2/X1X2Y condition. In Aspidoscelis deppii (Wiegmann, 1834) (2n = 50) and Anolis capito Peters, 1863 (2n = 42), we found a different karyotype from the ones previously reported for these species. Moreover, in A. capito , the cytogenetic observation is concurrent with a considerable genetic divergence (9%) at the studied mtDNA marker (MT-ND2), which is indicative of a putative new cryptic species. The skink Scincella cherriei (Cope, 1893), showed high values of genetic divergence (5.2% at 16S gene) between the specimens from Montes Azules and those from Costa Rica and Nicaragua, comparable to the values typical of sister species in skinks. A lower level of genetic divergence, compatible with an intraspecific phylogeographic structure, has been identified in Lepidophyma flavimaculatum Dumeril, 1851. These new data identify taxa that urgently require more in-depth taxonomic studies especially in these areas where habitat alteration is proceeding at an alarming rate.

We characterize the amphibian and reptile fauna from four habitats of the Andean region of the department of Santan­der, Colombia. Eight species were recorded, including four amphibians and four reptiles. The family Hylidae was represented by two species and had the greatest abundance of individuals at 241. Among reptiles, the family with the greatest diversity was the Colubridae, with two species, but the Dactyloidae had the greatest abundance, with seven individuals. The grassland habitat exhibited the greatest diversity, with five species found, and it was the habitat with the greatest abundance of individuals, with 242 individuals obtained. The habitat with the least diversity and abundance was cropland, which had no species recorded.&nbsp

The mosquito Culiseta melanura (Coquillett) is the primary enzootic vector of eastern equine encephalitis virus (EEEV), a zoonotic Alphavirus endemic to eastern North America. In its northern range, Cs. melanura is considered a strict avian biter, transmitting EEEV among susceptible birds in a cycle of enzootic amplification. In its southern range, however, Cs. melanura is more general in host use, feeding heavily upon birds but also reptiles and mammals. The goal of this study was to better understand how host use of Cs. melanura changes throughout the year in Florida, where year-round EEEV transmission is observed. Mosquitoes were sampled in 2018 from nine sites across three central Florida counties. In total, 213 Cs. melanura bloodmeals were identified by PCR consisting of 39 species of birds, reptiles, and mammals. Avian bloodmeals were prominent throughout the year (range = 30–85%), and songbirds were a large portion of identified bloodmeals (37.1%). Reptiles surpassed birds only in spring (April–June), and brown anole (Anolis sagrei Duméril and Bibron, 1837 [Reptilia: Dactyloidae]) was the most commonly detected single host species (22.1% overall). Mammalian bloodmeals were mainly observed in summer, with humans being the most fed on mammal (12.7% overall). This study reveals that in southern foci of EEEV transmission, Cs. melanura host use varies throughout the year with reptiles providing the majority of blood meals in spring (51.3%), and birds are fed on more than other host groups during all other seasons (50.6–70.1%). In addition, feeding on mammals increases during summer months, which may implicate Cs. melanura in epizootic transmission in Florida.

Helminthological studies may contribute with valuable information on host biology and conservation. Herein, we provide new data on helminths infecting the lizard Norops fuscoauratus, testing one of the factors considered most important in parasitic ecology: host size. We analysed 25 specimens of N. fuscoauratus from three highland marshes in the Brazilian semi-arid. Eight taxa of helminths belonging to Nematoda, Trematoda and Acanthocephala were found. Physaloptera sp. showed the higher prevalence (40%), with a mean intensity of infection of 3.3 ± 1.46 (1–16) and mean abundance 1.32 ± 0.65 (0–16). Norops fuscoauratus represents four new host records for the helminths Cyrtosomum sp., Pharyngodon travassosi, Strongyloides sp. and Centrorhynchus sp. There is no relationship of host body size (P = 0.79) and mass (P = 0.50) with parasite richness. In addition, the present study contributes to the knowledge of the parasitic fauna of N. fuscoauratus and the Neotropical region.

The ecomorph concept of the adaptive radiation of Caribbean anoles is characterized by a suite of behavioral, ecological, and morphological traits that are tightly linked to microhabitat use in lizards. However, most studies on the adaptive radiation of anoles have been conducted in a single macrohabitat type—lowland tropical forests. Because behavior can help organisms cope with different environmental conditions, we can predict that there will be key shifts in behavior within ecomorphs when examined across different macrohabitats, although this idea remains empirically underexplored. Here we utilized the replicated evolution of montane endemics from a primarily lowland species in a clade of trunk–ground Anolis lizards to test the hypothesis that shifts in basking behavior, wariness, and display behavior accompany divergence into montane habitats. The montane specialists A. armouri and A. shrevei each independently evolved from the primarily lowland dwelling A. cybotes in two widely separated mountain chains on the island of Hispaniola. We found evidence for a convergent behavioral response to the high-altitude macrohabitat: A. armouri and A. shrevei spend more time basking, utilize more open environments, and are warier than lowland A. cybotes. We also found divergence in display behavior in A. shrevei. We detected no evidence of divergence in locomotor behavior with elevation among active lizards. Together, our results suggest that the ecomorph concept would be enriched by extending observations of behavior (and other aspects of the phenotype) into different macrohabitats. Future work should focus on whether the observed behavioral shifts are clinal, reflecting local adaptation within A. cybotes, or fixed differences between the lowland generalist and montane species. Adaptation to the macrohabitat has previously been underappreciated as a source of behavioral diversity in Anolis lizards; this study is the first step toward documenting intraecomorph behavioral variation across divergent habitats.

Reproduction was studied in an invasive population of Anolis carolinensis in the Hawaiian Islands, USA. Timing of events in the reproductive cycle was similar between A. carolinensis populations in Hawaii and native populations of the species in the southeastern United States. In Hawaii, males of A. carolinensis undergo a prolonged period of spermiogenesis (sperm formation) starting in November (n=1) and December (n=1) and continuing into August. Gravid A. carolinensis females in Hawaii (n=40) produce one egg in continuous succession from March into August. Reproductive activity in A. carolinensis in Hawaii ceased prior to the colder, wetter, winter months.

We update the southeastern distribution of Anolis fuscoauratus d’Orbigny, 1837 based on newly collected specimensand on specimens from scientific collections misidentified or deposited with questionable identifications. We includephotographs showing the coloration of live individuals and a distribution map for this species in the southeasternAtlantic Forest of Brazil. Although this species is rather rare, it is more regionally widespread than previously realizedand has an altitudinal range of approximately 30–800 m above sea level.

The predation of a free-living Tropical House Gecko Hemidactylus mabouia (Moreau de Jonnes, 1818) by the Spotted Brown Trope, Tropidophis pardalis (Gundlach, 1840), is recorded for the first time. Other two vertebrates, the Cuban Green Anole, Anolis porcatus Gray, 1841 (Squamata: Dactyloidae) and the small frog Eleutherodactylus planirostris (Cope, 1862) are also recorded as preys of this dwarf boa in the same locality.

This article present the case of an adult female of the katydid Erechthis gundlachi Bolívar, 1888 (Orthoptera: Tettigoniidae: Conocephalinae: Agraeciini) eating a very young juvenile of the lizard Anolis homolechis (Cope, 1864).

The salticid spider Phidippus regius C.L. Koch, 1846 is documented preying on small frogs (Hyla spp., Osteopilus septentrionalis) and lizards (Anolis carolinensis and Anolis sagrei) in Florida, USA. Female as well as male P. regius were engaged in feeding on this type of vertebrate prey. A total of eight incidents of P. regius devouring vertebrates have been witnessed in seven Florida counties. Furthermore, we report an incident of a large unidentified Phidippus sp. (possibly P. bidentatus F. O. Pickard-Cambridge, 1901) preying on an immature anole lizard in Costa Rica. P. regius, otherwise known to feed almost exclusively on insects and spiders, is one of the world's largest salticid spiders reaching a maximum recorded body length of 2.2 cm. Most other salticid spiders appear to be too small in body size to overcome vertebrate prey. Vertebrate predation by salticid spiders has not been previously documented in the scientific literature. Together with Salticidae, spiders from 27 of 114 families (24%) are currently known to occasionally consume vertebrate prey.

Anoline lamellae terminate in an epidermal free margin carrying the majority of its setae. How the free margin is extruded from the body of the scale is not well understood. Two hypotheses have been advanced to account for it, one advocating distal migration of the outer epidermal layers relative to the body of the lamella, and the other proposing regression of the dermal core. Available evidence provides partial support for both. We assembled a series of specimens of Anolis grahami representing all shedding cycle stages, and prepared histological sections of the toe pads to allow measurement of appropriate lamellar components through the shedding cycle. Through its proliferative phases the lamellae increase markedly in length, with the distance between the distal tip of the dermal core and that of the lamella accounting for most of this, indicating that epidermal extrusion is responsible for production of the new free margin. The dermal core showed no evidence of regression. Concomitant with epidermal extrusion, the lacunar cells on the inner lamellar face hypertrophy and keep pace with the increasing thickness of the outer lamellar face resulting from the lengthening of the replacement setae. The integrated changes observed are consistent with continuity of functioning and alignment of the exposed setal carpet of the outer epidermal generation while ensuring that the new setal carpet is fully aligned and functional immediately after shedding. At shedding the original proportions of the lamellae are restored. Development of the new free margin results from a combination of distal displacement of Oberhautchen cells along with arrested maturation of the epidermis in this region. Changes in length of the lamellae during the proliferative stages may impact the overall size of the adhesive toe pad, which may have consequences for assessments of the relationship between whole animal clinging ability and adhesive pad area. J. Morphol. 278:360-368, 2017. © 2017 Wiley Periodicals, Inc.

The eminent Austrian zoologist Franz Werner described several new species of amphibians and reptiles from America, including Anolis aequatorialis Werner, 1894 and Hylodes appendiculatus Werner, 1894. Both species were described based on single specimens, with no more specific type localities than “Ecuador” (Werner 1894a,b). After its description, A. aequatorialis remained unreported until Peters (1967) and Fitch et al. (1976) published information on its distribution and natural history. Anolis aequatorialis is currently known to inhabit low montane and cloud forest on the western slopes of the Andes from extreme southern Colombia to central Ecuador, between 1300 and 2300 m elevation (Ayala-Varela & Velasco 2010; Ayala-Varela et al. 2014; Lynch et al. 2014; D.F. Cisneros-Heredia pers. obs.). Likewise, Hylodes appendiculatus (now Pristimantis appendiculatus) remained only known from its type description until Lynch (1971) and Miyata (1980) provided certain localities and information on its natural history. Pristimantis appendiculatus is currently known to occur in low montane, cloud, and high montane forests on the western slopes of the Andes from extreme southern Colombia to northern Ecuador between 1460 and 2800 m elevation (Lynch 1971; Miyata 1980; Lynch & Burrowes 1990; Lynch & Duellman 1997; Frost 2016). To this date, the type localities of both species remain obscure. The purpose of this paper is to restrict the type localities of Hylodes appendiculatus Werner, 1894 and Anolis aequatorialis Werner, 1894 based on analyses of the travel journals of their original collector.

Iguanians (Pleurodonta) are one of the reptile lineages that, like birds and mammals, have sex chromosomes of ancient origin. In most iguanians these are microchromosomes, making a distinction between the X and Y as well as between homeologous sex chromosomes in other species difficult. Meiotic chromosome analysis may be used to elucidate their differentiation, because meiotic prophase chromosomes are longer and less condensed than metaphase chromosomes, and the homologues are paired with each other, revealing minor heteromorphisms. Using electron and fluorescent microscopy of surface spread synaptonemal complexes (SCs) and immunolocalization of the proteins of the SC (SYCP3), the centromere, and recombination nodules (MLH1), we examined sex chromosome synapsis and recombination in 2 species of anoles (Dactyloidae), Anolis carolinensis and Deiroptyx coelestinus, in which the sex chromosomes represent the ancestral condition of iguanians. We detected clear differences in size between the anole X and Y microchromosomes and found an interspecies difference in the localization of the pseudoautosomal region. Our results show that the apparent homomorphy of certain reptile sex chromosome systems can hide a cryptic differentiation, which potentially may influence the evolution of sexual dimorphism and speciation.

The egg tooth is a hatching adaptation, characteristic of all squamates. In brown anole embryos, the first tooth that starts differentiating is the egg tooth. It develops from a single tooth germ and, similar to the regular dentition of all the other vertebrates, the differentiating egg tooth of the brown anole passes through classic morphological and developmental stages named according to the shape of the dental epithelium: epithelial thickening, dental lamina, tooth bud, cap and bell stages. The differentiating egg tooth consists of three parts: the enamel organ, hard tissues and dental pulp. Shortly before hatching, the egg tooth connects with the premaxilla. Attachment tissue of the egg tooth does not undergo mineralization, which makes it different from the other teeth of most squamates. After hatching, odontoclasts are involved in resorption of the egg tooth's remains. This study shows that the brown anole egg tooth does not completely conform to previous reports describing iguanomorph egg teeth and reveals a need to investigate its development in the context of squamate phylogeny.