Anolis lososi, Torres-Carvajal & Ayala-Varela & Lobos & Poe & Narváez, 2017

Torres-Carvajal, Omar, Ayala-Varela, Fernando P., Lobos, Simón E., Poe, Steven & Narváez, Andrea E., 2017, Two new Andean species of Anolis lizard (Iguanidae: Dactyloinae) from southern Ecuador, Journal of Natural History 52 (13 - 16), pp. 1067-1089 : 1078-1086

Anolis lososi sp. nov.

( Figures 5 , 6 )

Proposed standard English name: Losos’ s anole

Proposed standard Spanish name: Anolis de Losos Holotype

QCAZ 10173 ( Figures 5 , 6 ), adult male, Ecuador, Provincia Zamora Chinchipe, Romerillos Alto , 4.227°S, 78.939°W, WGS84 , 1550 m, 18 December 2009, collected by Steven Poe, Levi Gray, Julian Davis, and Fernando Ayala .

Paratypes (23)

ECUADOR: Provincia Zamora Chinchipe: QCAZ 6850 , San Francisco Research Station, 3.971°S, 79.078°W, 1970 m, 2 April 2005, collected by Kristin Roos, Alban Pfeiffer, Andy Fries, Ulf Soltau and Florian Werner; QCAZ 10171–72 , road from Zamora to Loja, 3.970°S, 79.063°W, 1706 m, 17 December 2009, collected by Steven Poe, Levi Gray, Julian Davis and Fernando Ayala; QCAZ 10174–75 , same collection data as holotype; QCAZ 14125 , San Francisco Research Station , 3.973°S, 79.077°W, 1872 m, 9 September 2015, collected by Andrea Narváez and Leonardo Cedeño; QCAZ 14126 , San Francisco Research Station , 3.974°S, 79.078°W, 1883 m, 10 September 2015, collected by Andrea Narváez and Leonardo Cedeño; QCAZ 14128 , San Francisco Research Station , 3.973°S, 79.078°W, 1821 m, 11 September 2015, collected by Andrea Narváez and Leonardo Cedeño; QCAZ 14129 , San Francisco Research Station , 3.972°S, 79.077°W, 1912 m, 11 September 2015, collected by Andrea Narváez and Leonardo Cedeño; QCAZ 14130 , San Francisco Research Station , 3.972°S, 79.079°W, 1883 m, 11 September 2015, collected by Andrea Narváez and Leonardo Cedeño; QCAZ 14132 , San Francisco Research Station , 3.973°S, 79.078°W, 1857 m, 15 September 2015, collected by Andrea Narváez and Leonardo Cedeño; QCAZ 14133 , San Francisco Research Station , 3.973°S, 79.077°W, 1878 m, 15 September 2015, collected by Andrea Narváez and Leonardo Cedeño; QCAZ 14335–336 , San Francisco Research Station , canal path, 3.972°S, 79.075°W, 1898 m, 12 January 2016, collected by Omar Torres – Carvajal, Diego A . Paucar and reptile class students; QCAZ 14337 , San Francisco Research Station, Tarabita’ s path, 3.972°S, 79.079° W, 1848 m, 14 January 2016, collected by Fernando Ayala, Steven Poe and Chris Anderson; QCAZ 14434–439 , San Francisco Research Station , 3.971°S, 79.079°W, 1789 m, 12 January 2016, collected by Fernando Ayala, Steven Poe and Chris Anderson; QCAZ 14611 , San Francisco Research Station , 3.973°S, 79.077°W, 1872 m, 14 September 2015, collected by Andrea Narváez and Leonardo Cedeño; FHGO 1756 , 3.6 km southwest from La Pituca, 4.156°S, 78.974°W, 1820 m, 24 April 1998, collected by Diego Almeida- Reinoso and Fernando Nogales-Sornosa .

Diagnosis

Anolis lososi differs from other species of Anolis (clade Dactyloa ) from Ecuador and Peru, except Anolis orcesi , Anolis peruensis and Anolis williamsmittermeierorum , in having large smooth head scales, homogeneous lateral lepidosis, short limbs, short tail, and in lacking a rostral proboscis. Anolis lososi differs from A. orcesi , A. peruensis and A. williamsmittermeierorum in having differently coloured male and female dewlaps ( A. orcesi male: yellow with greenish blue at base, female: orange; A. peruensis male: solid yellow, female: black; A. williamsmittermeierorum male: tan with peach-orange distal edge, female: black and white; A. lososi male: white, female: black and orange). Anolis lososi , A. peruensis and A. williamsmittermeierorum lack a vertebral crest (present in A. orcesi ). Anolis lososi and A. williamsmittermeierorum have a black throat with bright yellow-orange mouth edges (grey in A. peruensis and black in A. orcesi ).

Anolis lososi is most similar morphologically to A. williamsmittermeierorum in lepidosis. It can be further distinguished from A. williamsmittermeierorum in having a longer dewlap fold and more lamellae under phalanges III–IV of fourth toe ( Table 3).

Finally, although the ND2 genetic distance between A. lososi and its closest relative A. williamsmittermeierorum is relatively low (0.108), it is comparable to DNA divergences between other inarguable species pairs, such as Anolis microtus versus Anolis insignis (0.100) and Anolis peraccae versus Anolis anchicayae (0.107) ( Figure 7 ; Supplemental material).

Description of holotype (scores for paratypes in parentheses)

SVL 61.4 mm (55.1−59.8 mm); tail length 77.2 mm (66.5−78.2 mm); head length 16.7 mm (14.0− 16.7 mm); head width 8.5 mm (7.8−9.4 mm); head height 7.4 mm (7.1−7.8 mm); humerus length 8.4 mm (7.8−9.5 mm); ulna length 7.9 mm (5.3−7.9 mm); femur length 10.3 mm (9.6−10.6 mm); tibia length 9.7 mm (7.6−9.7 mm); dewlap length 29.1 mm (20.8 −29.1 mm); dewlap height 13.4 mm (6.8−13.4 mm); interparietal length 2.2 mm (1.8 −2.3 mm); ear opening vertical length 1.5 mm (1.3−1.5 mm); snout length 7.7 mm (7.0 − 7.7 mm); interorbital distance 2.1 (2.1−2.5 mm).

Head scales smooth (smooth to rugose); 4 (4 − 6) scales between second canthals; 6 (3 − 6) scales bordering the rostral posteriorly; circumnasal in contact with rostral (in contact or separated by one scale); supraorbital semicircles in contact; supraocular disc with 3 (3 − 5) abruptly enlarged scales; one short and rectangular superciliary followed by quadrangular scales (or small scales); 2 (2 − 3) loreal rows; 5 (5–15) loreal scales; interparietal smaller (smaller to similarly sized) than ear opening, in contact (separated by two small scales) with semicircles anteriorly; suboculars in contact with supralabials; 7 (6 − 8) supralabials counted up to a point below centre of eye; 2 (2 − 4) postmentals; 5 (2 − 5) enlarged sublabials in contact with infralabials.

Low nuchal crest formed by conical scales; low body crest formed by conical scales alternating with smooth scales (crest present in adults of both sexes); dorsal scales smooth (smooth or swollen); 10 (7 − 9) mid-dorsal scales in a longitudinal segment representing 5% of SVL; flank scales more or less barely separated by skin; ventral scales similar in size to dorsals (ventrals larger than dorsals or similarly sized), smooth,

imbricate (juxtaposed or subimbricate) and arranged in transverse rows; 8 (6 − 8) midventral scales in a longitudinal segment representing 5% of SVL.

Toepads overlapping the first phalanx in all toes; 20 (18 − 20) lamellae under third and fourth phalanges of fourth toe; supradigitals smooth; tail weakly compressed, with a single row of mid-dorsal scales forming a serrate crest; enlarged postcloacal scales present (absent in females).

Nuchal and dorsal folds absent in both sexes; dewlap large, extending posteriorly behind fore limbs (same but slightly smaller in females), with six longitudinal single rows of scales similar in size to ventrals, separated by naked skin.

Variation in morphological characters in A. lososi is presented in Table 2.

Colour in life

Holotype ( Figure 6 (a,b); adapted from colour photos of stressed specimen): head, body, limbs and tail with brown, yellow and green marks giving a lichenous appearance; supralabials and infralabials with a dark brown speck each; light band extending from tympanum to axilla; body with four broad transverse dark bands from shoulder to sacrum, as well as reticulating brown lines and small black spots mid-dorsally; ventral surface of head, body, limbs and tail cream with a few brown dots; limbs with narrow transverse bands formed by yellow dots; tail with dark brown transverse bands; iris almond brown; dewlap skin and scales cream.

Adult male QCAZ 10171 ( Figure 6 (c,d), adapted from colour photos): general colour pattern similar but lighter than holotype; edge of mouth including jaw hinges bright orange; tongue russet brown. Several male specimens had pale blue apicogorgetal and apicosternal regions.

Adult female QCAZ 10175 ( Figure 5 (e,f); adapted from colour photos): head, body, limbs and tail pale dark tan (pale green lichenous appearance); head flanks with a blotch anteriorly to the tympanum light sap greenish grey; limbs with narrow transverse bands formed by dark olive dots; tail with dark brown transverse bands; body flanks with yellowish grey dots; ventral surface of head, body, limbs and tail whitish cream with dots greyish brown; dewlap skin brilliant orange-yellow with round black blotches, appearing jaguar-striped; dewlap scales cream; iris moderate orange.

Gular sac radiance

We obtained and averaged colour data from four females and eight males measured in vivo on the day of collection. In males, the gular sac reflected more or less equally all sections of the visible spectrum, featuring cream-white colours – reflectance increased from 400 nm to 500 nm (55%), after which it remained steady ( Figure 3 ). The centre exhibited a slight decrease on reflectance at high wavelength values. The base and region close to the head were brighter than the edge and the section close to the abdomen. In females, reflectance increased steadily with wavelength ( Figure 3 ); orangeyellow patches were measured at the base, edge and in areas closest to the head and abdomen, whereas dark patches were measured at the centre. Males exhibited more UV reflection (i.e. wavelength <400 nm) in all measured areas than females.

Distribution and ecology

Anolis lososi occurs on the Amazonian slopes of the Cordillera Real in southern Ecuador, Provincia Zamora Chinchipe, between 1550 and 1970 m ( Figure 4 ). It is known from the upper basin of the Zamora river (Atlantic drainage) in evergreen lower montane forest ( Homeier et al. 2008). This area has suffered from dramatic deforestation ( Tapia-Armijos et al. 2015). However, most individuals of A. lososi were collected within Podocarpus National Park, which suggests that at least some populations of this species are well protected.

Specimens of A. lososi were collected mainly in open areas. At the type locality, they were found along a secondary forest edge near a creek; at the San Francisco Research Station, they were collected along trails and in both primary and secondary forest gaps, and in disturbed areas. All individuals were found at night (19:00h and 24:00h) sleeping horizontally on twigs of bushes and trees, as well as on leaves of arboreal ferns between 2 and 8 m above ground. A male and female (QCAZ 10171–10172) were found together near a river (about 5 m from shore); they were 10 cm away from each other, 2 m above the ground, resting horizontally with their heads toward the distal end of the branch. This species occurs in sympatry with A. podocarpus and A. soinii at the type locality, and with A. soinii and A. hyacinthogularis at San Francisco Research Station.

Etymology

The specific epithet lososi is a noun in the genitive case and is a patronym for Jonathan B. Losos, who has dedicated his life to the study of anole lizards. After visiting Ecuador a few years ago, he inspired young Ecuadorian biology students who are undertaking pioneering studies on the ecology of these lizards.

Figures

Figure 3. Radiance of the gular sac in males and females of Anolis hyacinthogularis sp. nov. and Anolis lososi sp. nov. Measured regions are gular sac base ‘b’, centre ‘m’, edge ‘e’, as well as regions close to the head ‘h’ and close to the abdomen ‘a’. Photographs by A. Narváez and M. J. Quiroz.

Figure 4. Distribution of Anolis hyacinthogularis sp. nov. (left) and Anolis lososi sp. nov. (right) in Ecuador.

Figure 5. Head of the holotype (QCAZ 10173) of Anolis lososi sp. nov. in dorsal (top), lateral (middle), and ventral (bottom) views. Photographs by V. Chasiluisa. Scale bar = 5 mm.

Figure 6. Anolis lososi sp. nov. Holotype, adult male (SVL = 61.4 mm, QCAZ 10173, a, b), adult male (SVL = 55.1 mm, QCAZ 10171, c, d), subadult female (SVL = 45.0 mm, QCAZ 10175, e, f), and adult female (SVL = 59.8 mm, QCAZ 10172, g, h). Photographs by O. Torres-Carvajal (a, b, e, f); F. Ayala (c, d, g, h).

Figure 7. Phylogeny of Dactyloa including Anolis hyacinthogularis sp. nov. and Anolis lososi sp. nov. Maximum clade credibility tree obtained from a Bayesian analysis of 114 specimens, two mitochondrial genes (COI, ND2) and one nuclear gene (RAG1). Numbers above branches correspond to Bayesian posterior probability (PP) values; asterisks represent PP ≥ 0.95. GenBank accession numbers along with locality data are presented in Table 1 for all specimens included in this tree.