Phyllodactylus benedettii, Ramírez-Reyes & Flores-Villela, 2018

Ramírez-Reyes, Tonatiuh & Flores-Villela, Oscar, 2018, Taxonomic changes and description of two new species for the Phyllodactylus lanei complex (Gekkota: Phyllodactylidae) in Mexico, Zootaxa 4407 (2), pp. 151-190 : 168-174

Phyllodactylus benedettii sp. nov.

Holotype. Adult male ( MZFC 28774 ) collected at Chamela Biological Station , La Huerta, state of Jalisco (19.50 N, - 105.44W WGS84) on September 15, 2014 by Tonatiuh Ramírez Reyes and Alfredo Villarruel.

Paratypes (14). An adult female ( MZFC 28773) collected at Chamela Biology Station, La Huerta municipality (19.50 N, - 105.44W) on September 15, 2014 by Tonatiuh Ramírez Reyes and Alfredo Villarruel; ( MZFC 21817- male, MZFC 21818-male, MZFC 21877, MZFC 21878-male, MZFC 21879, MZFC 21880-female, MZFC 21885- male) collected at the Chamela Biology Station, La Huerta municipality (19.50 N, - 105.44 W) by Oscar Flores Villela in 2007; an adult female ( IBH 2133-2) and five adult males ( IBH 2133-3, 2133-4, 2133-5, 2133-7, 2133-9) collected at 2 km. SE, of the Station of Biology, Chamela (19.51 N, - 104.94 W) on October 27, 1971 by Cornelio Sánchez; an adult female ( IBH 2138-4) collected at 5 km. S., de Chamela, Biology Station, UNAM, municipality La Huerta (19.49 N, - 104.95 W) on May 25, 1974 by Gustavo Casas Andreu.

Etymology. The species is dedicated to the memory of the great Uruguayan writer Mario Benedetti in recognition of his prolific literary production and critical thinking of great importance in the political and social life of Latin America.

Diagnosis. Phyllodactylus benedettii sp. nov. is a species with the largest average size within the P. lanei complex, with a maximum SVL of 75.2 mm. Dixon (1964) established three fundamental characteristics to include species and subspecies within the P. lanei complex and differentiate them from P. tuberculosus : species of the P. lanei complex have a low number of interorbital scales (12–19), white venter and low number of scales across the snout between the third labials (does not specify range of values). Phyllodactylus benedettii shows white-yellow venter, 14.4 interorbital scales on average, and 22.3 scales across the snout between the third labials.

Within the P. lanei complex (including insular and continental species), the only species that exceed 70 mm of SVL are P. lupitae and P. lanei , two species that are very isolated geographically. It is clearly different from the nearest species P. rupinus , which has a maximum SVL of 69.4 mm, considered as a medium size species. According to the statistical analyzes previously carried out, the combination of diagnostic morphological characters are the following (in addition to the SVL): It differs clearly from all species of Phyllodactylus as it presents 62.6 (60–69) longitudinal scales, 27 (24–30) tubercles from head to tail, 14 (11–13) interorbital scales, 29 (25–32) scales across venter, 22 (22–23) third labial–snout scales, 13 (13–14) rows of tubercles across dorsum. Although P. benedettii is very similar to P. lupitae in some meristic characters, they present differences in the morphometric space ( Fig. 7 ). Some measures allow us to differentiate these species are the head-length and axillagroin length: P. benedettii (17.9 mm and 28.4 mm) and P. lupitae (16.86 mm and 27.93 mm). According to Castiglia et al. (2009; 2010), the karyotypes of P. l. lanei (Tierra Colorada, Guerrero) and P. l. rupinus from Chamela, Jalisco (here named P. benedetti ) are different. According to their studies P. lanei has 2n = 33–34 and FN = 40–41, while P. benedettii has 2n = 38 and FN = 38.

Phyllodactylus benedettii has fewer than 30 tubercles from head to tail (24–30, 27.4 avg.) similar to P. lupitae (27–31, 28.8 avg.), P. paucituberculatus (28.7), P. kropotkini (25–31, 28.4 avg.) and P. rupinus (26–30, 28 avg.); other species have more than 30 tubercles from head to tail on average: P. lanei (32–34, 33.2 avg.), P. isabelae (30– 35, 32.2 avg.), P. t. magnus (35–40, 37 avg.), P. muralis (30–40, 33 avg.) and P. tuberculosus (33–41, 36.4 avg.). Phyllodactylus benedettii showed 62 longitudinal ventral scales (LVS) on average (60–69), other values were P. isabelae (52–58, 56.2 avg.), P. lupitae (60–64, 61.8 avg.), P. lanei (60–74, 66 avg.), P. rupinus (56–67, 63 avg.), P. kropotkini (63–73, 67 avg.), P. t. magnus (52–58, 54 avg.), P. muralis (57–61, 59 avg.) and P. tuberculosus (51–64, 57 avg.). Phyllodactylus benedettii presents 14.4 interorbital scales similar to P. kropotkini (14–16, 14.6 avg.) and differs from P. isabelae (14–20, 15.1 avg.), P. lupitae (14–20, 16.8 avg.), P. lanei (15–17, 15.5 avg.), P. rupinus (14–17, 15.5 avg.), P. t. magnus (21–23, 23.3 avg.), P. muralis (21–27, 23.2 avg.), and P. tuberculosus (16–22, 19.2 avg.). Phyllodactylus benedettii presents 29 scales across venter and differs from P. isabelae (26–29, 27.8 avg.), P. lupitae (26–29, 24.6 avg.), P. lanei (29–32, 30 avg.), P. rupinus (24–28, 26.5 avg.), P. kropotkini (29–33, 30.6 avg.), P. t. magnus (26–29, 27.3 avg.), P. muralis (33) and P. tuberculosus (27–33, 30.2 avg.). Phyllodactylus benedettii presents 22 third labial–snout scales on average (22–23) and differs from P. isabelae (19–24, 21.4 avg.), P. lupitae (23–28, 25.5 avg.), P. lanei (20–23, 21.2 avg.), P. rupinus (18–22, 20.6 avg.), P. kropotkini (19–21, 20.2 avg.), P. t. magnus (24–26, 24.6 avg.), P. muralis (24–28, 25.2 avg.) and P. tuberculosus (21–26, 24.2 avg.). Phyllodactylus benedettii presents 13.8 rows of tubercles across dorsum on average (13–14) and differs from P. isabelae (15–18, 16.7 avg.), P. lupitae (14–15, 14.8 avg.), P. lanei (14–16, 15.6 avg.), P. rupinus (13–15, 14 avg.), P. kropotkini (12– 14, 13.4 avg.), P. t. magnus (13–15, 14.3 avg.), P. muralis (12–13, 12.2 avg.) and P. tuberculosus (12–17, 14 avg.). Finally, P. benedettii presents the largest values in the following measures: 5.50 mm in length of the 4th toepad and 35.8 mm in axilla-groin length (LAG); compared to P. isabelae (4.1 mm, 22.2 mm), P. lupitae (4.9 mm, 31 mm), P. lanei (4.6 mm, 21.1 mm), P. rupinus (5 mm, 25.6 mm) and P. kropotkini (5 mm, 23 mm).

Description of holotype ( Fig. 9 ). (All bilateral counts are given as right/left). Adult male, robust body, head not flattened, neck slightly differentiated from the head. The head scales are granular, most are slightly globose, and are interspersed with circular-oblique flat scales that are mainly located between the eyes towards the tip of the snout. The rostral scale presents an intermediate striation near to middle of the scale (a groove that extends longitudinally to the middle part of the rostral scale); nostril bordered by a simple supranasal (one supranasal scale only), first labial, rostral and two postnasals; two supranasal scales contact each other, bordered by 7 postnasals; 22 interorbital scales beginning at the anterior ocular border; 15 scales across the snout from the second right labial, 24 scales across the snout at the level of the third labial scale on the right side; 8/9 loreal scales from the first ocular scale to the nostril; 11/10 supralabial scales; posterior border of the first supralabial in contact with the nostril (right and left); 8/8 infralabial scales; elongated auricular opening smaller than ocular diameter; occipital scales different in size and shape to the interorbital scales, while the interorbital scales are more or less uniform in size and shape (oval shape), they are smaller than the occipital ones. The occipital scales are larger than the interorbital scales, they have irregular shapes and presents some circular tubercles intermixed; mental scale is equally wide as long (3.6 mm), forms an irregular "V" towards the posterior section, two postmental scales closely in contact each other and with the first and second infralabials; postmental scales in contact with a row of 4 scales, the 2 intermediates larger than the extremes, 10 scales of uniform size border the row of scales.

Body with granular scales; 13 rows of strongly keeled dorsal tubercles with variable size; 25 tubercles in a middle dorsal row from base of the head to base of the tail; 21/15 tubercles from axilla to groin; 34 rows of cycloid scales across the venter; 87 longitudinal ventral scales from the first differentiated scale on the gular region to the cloacal opening; ventral scales larger compared to the lateral scales of the body and the gular region; scales slightly imbricate on the anterior and posterior extremities, scales dorsally granular intercalated with strongly keeled tubercles of variable size. The tail of the specimen is original (not regenerated) and measures 55.5 mm with the tip broken. Dorsal scales of the tail are imbricate with intercalated tubercles and juxtaposed ventral scales, keeled tubercles extend to more than half of the tail, as they extend to the tip of the tail the tubercles gradually flatten until they form keeled scales. Does not present femoral or precloacal pores.

Digital lamella formulae: right posterior (7-9-12-13-10), left posterior (6-8-11-13-9); right anterior (6-9-12-11- 11), left anterior (6-10-13-11-12), fourth finger of the extremities longer than others; digital toepads longer than wide on all fingers.

Measurements in mm: snout-vent length 70.9; axilla-groin 35.8; head length 16.5; head width 13.2; snout length (to eye) 10.05; eye diameter 4.1; auricular opening (maximum) 1.6; length of fourth finger 5.5; length of fourth toe 5.7; width between eye supercilliaries 9; internaral 2.1.

Coloration of holotype in ethanol ( Fig. 8 ). The holotype exhibits relatively little coloration, the dorsal background color varies from brown to creamy yellow, with dark brown faint irregular spots (less evident compared to live coloration), the ventral surface varies yellowish and cream; half of the ventral color on the tail to the tip is cream.

Pattern and color in life ( Fig. 10 ). Dorsally presents a clear background, a mixture of cream-pink, on this background, patterns of irregular patches dark brown color and some completely black spots are present, some of these spots are slightly surrounded by yellowish scales; on the middle dorsal part a longitudinal line of the background color runs from head to tail; scales around the eyes and some supralabials are slightly colored light yellow; ventrally presents a white-cream coloration in combination with light yellow, more evident towards the ventral region; the pupils are black on a bronze iris. Dorsal coloration of the tail is white-cream with brown stripes and some black points.

Variation. All meristic and morphometric characters are presented with mean values, standard deviation and range of meristic ( Table 2) and morphometric variables ( Table 7).

Distribution and habitat. Phyllodactylus benedettii sp. nov. is an endemic species to the state of Jalisco, restricted to the Chamela-Cuixmala Biosphere Reserve. According to the INEGI digital map of elevation (resolution of 15 m), the type locality is located at 100 masl, located in the biogeographic province of the Western Pacific surrounded by deciduous tropical forests ( Fig. 11 ). The microhabitats reported for this species are diverse: soil, trees, shrubs and in habitats modified by man as houses or bridges or some other constructions (meteorological stations). They are particularly abundant in rocks and crevices and in people's houses or buildings where it feeds on insects and other invertebrates ( García & Ceballos 1994; Ponce-Campos & García 2007; Vitt & Caldwell 2014; pers. obs.). Males of P. benedettii have an extended period of reproduction (August–March) with two peaks, one in rainy season (August–October) and another one in dry season (November–March). Reproductive behavior (courtship and mating) begins in August which coincides with the beginning of rainy season. The females produce clutches of two eggs, with up to three clutches during the breeding season, with a peak of egg production from December to March coinciding with the dry season ( Ramírez-Sandoval et al. 2006). This species presents a distinct karyotype from P. lanei presumably due to Robertsonian fusions/fissions with 19 pairs of telocentric chromosomes (2n = 38, FN = 38) ( Castiglia et al. 2009). They are lizards of nocturnal habits and feed on insects, although to date there have been no studies on the composition of their diet ( García & Ceballos 1994; Vitt & Caldwell 2014). Representative species of tropical dry forest of Chamela-Cuixmala Biosphere Reserve are: Sciadodendrom excelsum, Brosimum alicastrum, Orbignya cohune and Tabebuia donell-smithi ( Ceballos et al. 1999). The deciduous tropical forests is near the mouth of the Cuixmala River and is characterized by the abundance of water in the rainy season, the climate is tropical (warm-humid), has an annual average temperature of 25 °C and the annual rainfall varies from 748–1000 mm ( García & Ceballos 1994; Ceballos et al. 1999). Other species of reptiles with nocturnal activity that inhabit the deciduous tropical forests are: Hemidactylus frenatus, Boa constrictor, Lampropeltis triangulum, Leptodeira maculata, Pseudoficimia frontalis, Senticolis triaspis, Tropidodipsas philippii ( Sibon philippi ), Tantilla bocourti, Thamnophis valida, Trimorphodon biscutatus, Micrurus distans, Agkistrodon bilineatus, and Crotalus basiliscus ( García & Ceballos 1994) .

Comments on conservation and threats. This species inhabits the Chamela-Cuixmala Biosphere Reserve decreed in 1993. There is a minor threat posed by the localities bordering the reserve, and according to the observations made by the first author, apparently there is a large population of P. benedettii geckos inside Chamela- Cuixmala Biology Station (UNAM). There are two main threats: the false beliefs of some people who consider this species as poisonous (local inhabitants), and the possible negative effects due to the presence of invasive gecko Hemidactylus frenatus , a species with a very large population that may exploit the same ecological niche. To date there is no specific study on niche overlap between H. frenatus and Phyllodactylus species in Mexico, however, its potential danger to local biodiversity has been documented, including interspecific negative interactions with native geckos from different areas of the world; one example of this is the displacement and reduction of six gecko species of the genus Nactus on the Mascarene Islands and Lepidodactylus lugubris throughout Pacific islands. In these cases H. frenatus has been observed stalking, lunging towards and biting at other geckos (Global Invasive Species Database 2015). Due to the above, and as a potential risk to local biodiversity in Mexico, it is classified as a high risk invasive species ( CONABIO 2016).

Phyllodactylus kroPotkini sp. nov.

Holotype. Adult male ( MZFC 28736 ) collected in Nueva Filadelfia ( Huerta Vieja ) in the municipality of Tlapehuala , Guerrero (18.29 N, - 100.49 W WGS84), collected on March 15, 2014 by Tonatiuh Ramírez Reyes and Joel Rosas Avila.

Paratypes (4). Four male adults ( MZFC 28735 , MZFC 28737 , MZFC 28738 , MZFC 28739 ) collected at the type locality, Nueva Filadelfia ( Huerta Vieja ), Tlapehuala , Guerrero (18.29 N, - 100.49 W WGS84) collected on March 16, 2014 by Tonatiuh Ramírez Reyes and Joel Rosas Ávila.

Etymology. The species is dedicated to the memory of the great Russian philosopher, geographer and naturalist Piotr Kropotkin, who made great scientific and theoretical contributions about mutual support and altruism in some animal populations (including human society).

Diagnosis. Phyllodactylus kropotkini sp. nov. is a medium-sized gecko within the P. lanei complex. Dixon (1964) established three fundamental characteristics to include species and subspecies within the P. lanei complex and differentiate them from P. tuberculosus , species of the P. lanei complex have a low number of interorbital scales (12–19), white venter and low number of scales across the snout between the third labials (does not specify range of values). Phyllodactylus kropotkini has 14.6 interorbital scales on average (14–15), white venter and 20.2 third labial–snout scales on average (19–21). Phyllodactylus kropotkini presents a maximum recorded SVL of 62.3 mm that clearly differs from the nearest species P. lanei (max. 71 mm) and it differs from other species: P. isabelae (max. 57.5), P. lupitae (max. 73.3 mm), P. rupinus (max. 69.4), P. benedettii (max. 74.2) and P. bordai (max. 58.9 mm). Phyllodactylus kropotkini sp. nov. shows the following combination of characters: 28 (25–31) tubercles from head to tail, 67 (63–73) longitudinal scales, 13 (12–14) rows of tubercles across dorsum, 30 (29–33) scales across venter, 14 (14–15) interorbital scales and 20 (19–21) third labial–snout scales. Phyllodactylus kropotkini has a partial overlap with P. lanei and P. isabelae on morphometric space ( Fig. 7 ), however P. kropotkini is clearly differentiated from P. lanei by maximum length (SVL) ( Fig. 6 ); it also presents the follow measures: axilla-groin length (22.54 mm), snout length (9.34 mm), auricular opening (1.78 mm), while P. lanei has 24.7 mm axilla-groin length, 10. 61 mm snout length and auricular opening (1.95 mm) and P. isabelae has 18.7 mm axilla-groin length, 8.18 mm snout length and auricular opening 1.51 mm.

P. kropotkini has a low number of interorbital scales (14–15, 14.6 avg.) similar to P. benedettii (13–16, 14.4 avg.), all other species of Phyllodactylus exceed 15 interorbital scales on average, P. isabelae (14–16, 15.1 avg.), P. lupitae (14–20, 16.83 avg.), P. lanei (15–17, 15.5 avg.), P. rupinus (14–17, 15.5 avg.), P. t. magnus (21–23, 22.3 avg.), P. muralis (21–27, 23.2 avg.), P. tuberculosus (16–22, 19.2 avg.). There are an average of 30.6 scales across the venter (29–33), unlike P. isabelae (26–29, 27.8 avg.), P. lupitae (23–26, 24.6 avg.), P. lanei (29–32, 30 avg.), P. rupinus (24–28, 26.5 avg.) and P. benedettii (25–32, 29.1 avg.). Phyllodactylus kropotkini presents 67.6 longitudinal scales from the gular region to the anus on average (63–73), compared to P. rupinus (63–73, 63.6 avg.), P. lanei (60–74, 66 avg.), P. benedettii (60–69, 62.6 avg.), P. t. magnus (52–58, 54.6 avg.), P. muralis (57–61, 59 avg.) and P. tuberculosus (51–64, 57.7 avg.). This species also has 28.4 tubercles from head to tail on average (25–31), different from P. rupinus (26–30; 28 avg.), P. lanei (32–34, 33.2 avg.), P. benedettii , (24–30, 27.4 avg.), P. isabelae (30–35, 32.2 avg.), P. lupitae (27–31, 28.8 avg.), P. t. magnu s (35–40, 38 avg.), P. muralis (30–40, 33.6 avg.) and P. tuberculosus (33–41, 36.4 avg.) tubercles. It also presents 13.4 rows of tubercles on average across the dorsum (12–14), different from P. rupinus (13–15, 14 avg.), P. benedettii (13–14, 14.4 avg.), P. lupitae (14–15, 14.8 avg.), P. lanei (14–16, 15.6 avg.), P. isabelae (15–18, 16.7 avg.), P. t. magnus (13–15, 14.33 avg.), P. muralis (12– 13, 12.2 avg.) and P. tuberculosus (12–17, 14 avg.). Finally P. kropotkini presents 20.2 third labial–snout scales (19–21), similar to P. rupinus (18–22, 20.6 avg.) and it differs from the other species of Phyllodactylus : P. isabelae (19–24, 21.4 avg.), P. lupitae (23–28, 25.5 avg.), P. lanei (20–23, 21.2 avg.), P. benedettii (22–23, 22.3 avg.), P. t. magnus (24–26, 24.6 avg.), P. muralis (24–28, 25.2 avg.) and P. tuberculosus (21–26, 24.2 avg.).

Description of holotype ( Fig. 12 ). Adult male, medium body proportions (non-robust), short neck, head differentiated from the body. The head scales are granular, mostly flattened and some slightly globose towards the anterior part of the eye openings. Rostral scales have an intermediate longitudinal groove that reaches the middle part of the scale; nasal orifice bordered by a simple supranasal, rostral and 2 postnasals; nasal orifice in contact with the first supralabial, this has a fold that resembles a 3rd postnasal scale; 2 supranasal scales in contact with each other, bordered by 6 postmental scales; 25 interorbital scales from the anterior ocular border; 18 scales across snout starting with the 2nd labial; 26 scales across snout starting with 3rd labial scales; 12/12 loreal scales from the nostril to eye; 12/12 supralabials; edge of 1 st supralabial in contact with the nostril (right and left) forming a groove near mental scale (right and left); 10/11 infralabial scales; auricular opening elongated and smaller than ocular diameter; occipital scales different in size and shape to interorbital with some interspersed circular tubercles; mental scale forms an irregular "V" towards the posterior region; two postmental scales in contact each other and with the 1 st and 2nd infralabial scales (right and left); postmental scales in contact with a row of 5 scales, the central scale (pentagon shape) is larger than pair of adjacent scales (right and left). Body with granular scales, has 13 rows of strongly keeled dorsal tubercles of variable size; 30 tubercles in a dorsal row starting in the base of the head to base of the tail; 24/19 tubercles axilla-groin; 32 rows of cycloid scales across venter; 81 ventral longitudinal scales from the first differentiated scale in the gular region to the anus; ventral scales larger than the lateral sclaes of the body and the gular region; scales slightly imbricate on the four extremities. The tail of the specimen is original (not regenerated) and measures 39.6 mm with the tip broken, dorsal scales of the tail are subimbricate and mostly flattened. Therefore, the tubercles are clearly defined. Does not present femoral or precloacal pores. Lamellar formulae: anterior right leg (8-10-12-13-11), anterior left leg (9-10-13-13-11), posterior right leg (8-11-14-15-13), posterior left leg (8-12-14-14-13).

Measurements in mm. Snout-vent length 62.2; axilla-groin 23; head length 16.9; head width 12.3; snout length (to eye) 7.5; eye diameter 3.3; auricular opening (maximum) 1.7; length of fourth finger 5; length of fourth toe 6; width between eye supercilliaries 8.1; internaral 1.5.

Coloration of holotype in ethanol ( Fig. 12 ). The holotype presents a coloration that mixes the cream and light gray coloration on dorsal background, with a few dorsal dark brown spots; the ventral coloration is white with a slight light yellowish on posterior legs. The ventral coloration of the tail tip is dark gray, as well as the tips of the toepads.

Variation. All characters are presented with mean values, standard deviation and range of meristic ( Table 2) and morphometric variables ( Table 7).

Distribution and habitat. Phyllodactylus kropotkini is endemic to the middle region of the Balsas Basin at locality Nueva Filadelfia (Huerta Vieja), municipality of Tlapehuala, Guerrero ( Fig. 13 ). This species was observed mainly on rocks and between cracks in caves near the type locality. The specimens were collected in the house of a person who allowed collection inside his home. Due to its presumably small distribution area and its new taxonomic status, no further information on its reproduction or ecology is known. According to the digital elevation model of INEGI (resolution 15 m), the locality is at 579 m of elevation in relict lowland tropical dry forest in a habitat clearly disturbed by agricultural and livestock activity. Annual temperature for the locality is of 27 °C, with maximum temperatures of 38.6 °C and annual precipitation of 1099 mm on average. The Balsas Basin contains a great diversity of species of the genus Bursera , in fact this biogeographic province is considered the likely center of diversification and endemism of Bursera ( Toledo 2003; Pineda-García et al. 2007). Other species of reptiles that inhabit this province are: Hemidactylus frenatus, Ctenosaura pectinata, Sceloporus horridus, Urosaurus bicarinatus, Drymarchon corais, Leptodeira maculata, Salvadora mexicana, Crotalus simus, Micrurus laticollaris ( Reyna-Álvarez et al. 2010).

Comments about conservation and threats. This species inhabits the middle region of Balsas Basin, and the type locality is a remnant tropical dry forest; the species inhabits an area highly disturbed by agricultural and livestock activities, however, this species is able to adapt effectively to modified habitats, inhabiting houses or other types of buildings (bridges, fences, etc.). The main threat to this species (and in general to all species of the P. lanei complex) are the myths and false beliefs of the people, who consider this species as dangerous or poisonous ( García & Ceballos 1994). This false belief has led some people to implement "eradication" campaigns of this species in houses or wherever geckos are seen.

Taxonomic changes. Although the purpose of the present study was not to re-describe the species of the P. lanei complex, we will mention some points that are important to consider for the previously described species P. lanei , P. rupinus , P. isabelae and P. lupitae . The original description of P. lanei can be consulted in Smith (1935), P. rupinus can be consulted in Dixon (1964) and P. isabelae and P. lupitae in Castro-Franco & Uribe-Peña (1992) . The last publication includes an identification key for P. lanei , P. rupinus , P. isabelae and P. lupitae and agrees mostly with the results presented here.

The original descriptions are extensive enough and can be reviewed for a detailed taxonomic identification, here we will only mention characters that can help us identify the taxa.

Figures

FIGURE 7. Scatter plot of 11 morphometric variables for six species of the P.lanei complex:P.benedettii (black, group 1), P. kropotkini (aqua blue, group 2), P. rupinus (brown, group 3), P. lanei (blue, group 4), P. lupitae (red, group 5), P. isabelae (green, group 6). The first two components are shown with the percentage of variance on the axes.

FIGURE 8. Scatter plot of 19 bioclimatic variables to four species: P. lanei (green, group 1), P. rupinus (orange, group 2), P. benedettii (Black, group 3) and P. kropotkini (Red dot, group 4). The first two components are shown with the percentage of variance on the axes.

FIGURE 9. Dorsal and ventral view of the holotype of Phyllodactylus benedettii sp. nov. (MZFC 28774). The black bars represent the scale (10 mm).

FIGURE 10. Phyllodactylus benedettii sp. nov. from the type locality (Chamela-Cuixmala Biosphere Reserve, Jalisco) in life (Photo by Israel Solano Zavaleta).

FIGURE 11. Deciduos tropical forests from the type locality (Chamela-Cuixmala Biosphere Reserve, Jalisco) habitat of Phyllodactylus benedettii sp. nov. (Photos by Tonatiuh Ramírez Reyes).

FIGURE 12. Dorsal and ventral view of the holotype of Phyllodactylus kropotkini sp. nov. (MZFC 28736). The black bars represent the scale (10 mm).

FIGURE 13. Residuals of Tropical dry Forest at the type locality (Nueva Filadelfia, Tlapehuala, Guerrero) habitat of Phyllodactylus kropotkini sp.nov. (Photo by Tonatiuh Ramírez Reyes).

FIGURE6. Box plot of SVLfor sixspecies ofPhyllodactylus, on thehorizontal axis:P.isabelae (1), P.lupitae (2),P.lanei(3), P. rupinus (4), P. benedettii (5), P. kropotkini (6), the vertical axis indicates the length in mm. The medians are indicated by horizontal lines inside the boxes, the black dots indicate outliers.