Phyllodactylus pachamama, Koch, Claudia, Flecks, Morris, Venegas, Pablo J., Bialke, Patrick, Valverde, Sebastian & Rödder, Dennis, 2016

Koch, Claudia, Flecks, Morris, Venegas, Pablo J., Bialke, Patrick, Valverde, Sebastian & Rödder, Dennis, 2016, Applying n-dimensional hypervolumes for species delimitation: unexpected molecular, morphological, and ecological diversity in the Leaf-Toed Gecko Phyllodactylus reissii Peters, 1862 (Squamata: Phyllodactylidae) from northern Peru, Zootaxa 4161 (1), pp. 41-80 : 49-53

Phyllodactylus pachamama sp. nov.

Phyllodactylus reissii— Koch, Venegas & Böhme, Salamandra 42(2/3): 148. — 2006

Phyllodactylus reissii— Venegas, Townsend, Koch & Böhme, Journal of Herpetology 42(2): 393. — 2008

Phyllodactylus reissii— Koch & Beraun, Check List 7(3): 272. — 2011

Phyllodactylus reissii— Aurich, Koch & Böhme, North-Western Journal of Zoology 7(2): 310. — 2011

Phyllodactylus reissii (partim) ― Torres- Carvajal, Barnes, Pozo- Andrade, Tapia & Nicholls, Journal of Biogeography 41: 1887. — 2014

Phyllodactylus reissii (partim) — Aurich, Koch & Böhme, Phyllomedusa 14(1): 55. — 2015

Holotype. An adult male ( ZFMK 90886 ) from Balsas , Province of Chachapoyas, Department of Amazonas, Peru (6°50’45.0’’S, 77°59’47.9’’W, 1271 m a.s.l.), collected on 26 April 2009 by A. Garcia Bravo and C. Koch.

Paratypes (24). An adult male ( ZFMK 85004) and an adult female ( ZFMK 85005) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6°48’26’’S, 77°59’29’’W, 1000 m a.s.l.), collected on 10 July 2005 by P. Venegas and C. Koch; three adult females ( CORBIDI 5700, 5701; ZFMK 90885) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6°51’ S, 78°01’ W, 896—900 m a.s.l.), collected on 15 April 2009 by A. Garcia Bravo and C. Koch; an adult male ( CORBIDI 7710) and a subadult male ( ZFMK 91727) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6°51’ S, 78°02’ W, 870—919 m a.s.l.), collected on 30 March 2010 by J. Aurich and L. Y. Echevarria Espinosa; an adult female ( ZFMK 91725) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6°50’56.7’’ S, 78°01’32.4’’ W, 895 m a.s.l.), collected on 15 April 2010 by J. Aurich and L. Y. Echevarria Espinosa; four adult males ( CORBIDI 7711, 7712; ZFMK 91724, 91726) and a subadult female ( ZFMK 91728) from Balsas, Province of Chachapoyas, Department of Amazonas, Peru (6°51’ S, 78°01’ W, 920 m a.s.l.), collected on 27—29 May 2010 by J. Aurich and C. Laandauro Sanabria; two adult males ( CORBIDI 5703; ZFMK 90890), two adult females ( CORBIDI 5704, 5705), and a juvenile ( ZFMK 90889) from Zapatalgo, Province of Utcubamba, Department of Amazonas, Peru (6°05’ S, 78°29’ W, 830—1030 m a.s.l.) collected on 6—8 December 2009 by A. Garcia Bravo and C. Koch; an adult male ( CORBIDI 5706) and a subadult female ( ZFMK 90892) from Zapatalgo, Province of Utcubamba, Department of Amazonas, Peru (6°04’ S, 78°30’ W, 907 m a.s.l.) collected on 10 December 2009 by A. Garcia Bravo and C. Koch; a juvenile ( ZFMK 90891) from Zapatalgo, Province of Utcubamba, Department of Amazonas, Peru (6°06’05.2’’ S, 78°29’54.5’’ W, 521 m a.s.l.) collected on 9 December 2009 by A. Garcia Bravo and C. Koch; two adult males ( ZFMK 90887, 90888) and an adult female ( CORBIDI 5702) from San Vicente /Pusac, Province of Bolívar, Department of La Libertad, Peru (6°59’ S, 77°55’ W, 1425—1432 m a.s.l.) collected on 22 April 2009 by A. Garcia Bravo and C. Koch.

Diagnosis and comparison. Phyllodactylus pachamama sp. nov. is a comparatively large species with a maximum SVL of 77 mm. Among species of Phyllodactylus found in mainland South America only P. delsolari , P. dixoni , P. reissii and P. v en t r al i s exceed 70 mm SVL. Thus, this species can be distinguished from all other species by its larger size. In lacking an enlarged postanal plate it further differs from P. thompsoni and by the absence of a preanal plate it can be distinguished from P. angustidigitus , P. gerrhopygus , and P. heterurus . By having 10—16 well-defined rows of enlarged, trihedral keeled tubercles, the new species differs from P. angustidigitus , P. gerrhopygus , P. heterurus (dorsal tubercles absent in all three species), P. delsolari , P. inaequalis (fewer than 10 poorly defined rows of small, smooth, round tubercles in both species), P. microphyllus (dorsal tubercular rows indistinct, composed of small flat, oval tubercles), and P. thompsoni (10). By having 43—56 paravertebral tubercles between rear of head and posterior edge of thigh it differs from P. interandinus (54—83), P. k of ord i (31—36), P. pumilus (37—43), P. sentosus (26—31), and P. thompsoni (36—40). Phyllodactylus pachamama can be distinguished from P. heterurus , P. kofordi , P. pumilus , and P. sentosus by lacking tubercles on the base of tail. By the presence of tubercles on the tibia this species can further be differentiated from P. angustidigitus , P. clinatus , P. delsolari , P. gerrhopygus , P. inaequalis , P. interandinus , P. lepidopygus , and P. microphyllus . By the absence of tubercles on the thigh it can be distinguished from P. d i x o ni, P. k o f o rd i, P. pu m i l u s, P. s e n t o s u s, and P. ventralis . By the absence of tubercles on the forearm it can be differentiated from P. dixoni , P. kofordi , P. sentosus , and P. ventralis . From P. angustidigitus , P. microphyllus , and P. sentosus the new species further differs by having large terminal lamellae. It can be distinguished by the number of lamellae under the fourth toe (14—17, mean 15.1) from P. dixoni (11—13, mean 11.3), P. inaequalis (10—12, mean 10.7), P. johnwrighti (7—14, mean 10.5), P. k o f o rd i (11—13, mean 11.5), P. leoni (9—13, mean 10.9), P. lepidopygus (10—14, mean 11.5), and P. pumilus (11—13, mean 12.0). 40 % of specimens have 3 postmentals compared to only 1.9 % in P. reissii . In contrast to P. johnwrighti and P. pumilus (dark stripe from nostril to arm insertion always present), the postnasal and postorbital brown stripes in P. pachamama is usually absent or, if present, poorly defined.

Description of holotype ( Fig. 5 ). Head distinct from neck, covered with small granules that are largest on snout and smallest on interparietal region; rostral 3.3 mm wide, 2 mm high, somewhat rectangular, with a median groove along its posterior half; two internasals, in broad medial contact, each bordered posteriorly by two granules and a postnasal; nostril surrounded by rostral, first supralabial, two postnasals and internasal; 11 loreals between postnasal and anterior edge of orbit; scales in loreal region about three times larger than scales in interorbital region; 24 scales across snout at level of third supralabial, 17 at level of second supralabial; 12 supralabials, six to a point below the center of eye; eyes large, pupil vertically elliptical, with corrugated edges; orbital diameter 4.6 mm; eyelid with three visible rows of granules on left side and five on right side, granules of median row pointed in posterior part; 25 midorbital granules; interorbital distance 1.8 mm; 29 granules between posterior edge of eye and anterior edge of ear, eye to ear distance 7.1 mm; ear opening elliptical, posterodorsally oriented, 2.4 mm high and 0.7 mm wide, its anterior and posterior edges denticulated; rear of head intermixed with slightly larger tubercles; mental bell-shaped, larger than infralabials, 4 mm long, and 4 mm wide at its widest point; two postmentals, in contact with first pair of infralabials laterally and five scales posteriorly; nine infralabials, six to a point below center of eye, first two pairs largest, adjacent pairs of infralabials descreasing in size posteriard; scales on chin largest and juxtaposed in anterior part, decreasing in size and becoming more imbricate towards gular region.

Body depressed; dorsal surface of neck with small granules that are slightly smaller than those on the head; dorsum covered with small granules and 10 to 12 longitudinal rows of enlarged, trihedral keeled tubercles at midbody; six rows of tubercles from occiput to base of tail; paravertebral row with 44 tubercles between occiput and cloaca, 32 between axilla and groin; paravertebral rows separated from each other by about six granules; tubercles in paravertebral row separated from each other by 1—2 granules; 2—4 granules between other longitudinal tubercular rows; 110 scales around the midbody; ventral surface with smooth, imbricate scales, 74 from posterior gular region to cloaca, 26 transversely across midbody; dorsal surface of forearm with flattened, imbricate scales and without tubercles; thigh with flat, imbricate scales, largest in size anterodorsally and smallest posterodorsally, without tubercles; dorsal surface of tibia with a few scattered, hardly visible, cone-shaped or pulvinate tubercles, which are much smaller than the large tubercles on the back and only slightly larger than other scales on tibia; subdigital lamellae on left fingers (I—V) 8—10—12—13—11; subdigital lamellae on left toes (I—V) 8—11—15—15—13; terminal lamellae paired, greatly enlarged, truncated, the tip of the claw hardly visible between the two terminal pads, not projecting the terminal pads; tail complete, 1.18 times the SVL, covered with smooth and imbricate scales, on dorsal surface of tail about 3—4 times larger than dorsal body granules; medial subcaudals broadly enlarged, about two times as wide as long in anterior part of tail, decreasing in width posteriard.

Measurements in mm. — Snout-vent length 71; tail length 84; axilla-groin length 29; right forelimb length 24.0; right hindlimb length 31.9; head length 21.6; maximum head width 15.9; midorbital head width 12.3; head height 8.9.

Coloration of holotype in ethanol ( Fig. 5 ). Dorsal ground color of head, body and limbs pale brown, with irregular small brown flecks and blotches on the dorsal surface of the body and limbs; indistinct brown line extending laterally on each side from nostril through eye to above insertion of the forearm; lateral surface of the body cream with brown mottling on each scale; ventral surface of head and body cream, with brown mottling on each scale of the chest region and towards the venterolateral parts of head and body, scales on median part of venter and gular region almost without mottling; dorsal surface of tail with alternating cream, brown and dark brown blotches; subcaudal ground color cream, with dense grayish brown mottling; lamellae under fingers and toes grayish.

Variation (see also Table 3 ). SVL of male paratypes ranges from 48–75 mm (68.8 ± 4.3, n = 9), and SVL of female paratypes ranges from 44–77 mm (63.7 ± 11.6, n = 10). Tail length/total length is 0.43–0.56 (0.51 ± 0.03, n = 18). Male specimens possess 3—4 postanal tubercles on each side, which are slightly elevated. In females, postanal tubercles are lacking or only hardly visible and usually less pronounced as in males. The number of postmentals is 2—4 (2.5 ± 0.64). In one specimen (ZFMK 90887), postmentals contact first and second infralabials on each side, in all other specimens only first infralabial contacts postmentals, scales bordering postmentals 5—8 (6.4 ± 1). Two paratypes (ZFMK 90887, CORBIDI 5705) have 3 internasals, all others have 2 (2.1 ± 0.29). Lamellae on fourth finger 10—14 (12.5 ± 0.86); lamellae on fourth toe 14—17 (15.1 ± 0.81); supralabials 8—13 (10.2 ± 1.4), to a point below the center of eye 6—9 (7.7 ± 0.83); infralabials 7—11 (8.7 ± 1.03), to a point below the center of eye 5—7 (6.2 ± 0.7); loreals between postnasal and anterior edge of orbit 9—13 (11.6 ± 1.3); granules between posterior edge of eye and anterior edge of ear 24—28 (26.4 ± 1.1); scales across snout at level of third supralabial 22—26 (24.1 ± 1.4), at level of second supralabial 15—19 (17.1 ± 1.6); midorbital granules 23—25 (23.9 ± 0.86); longitudinal rows of dorsal tubercles 10—16 (13.2 ± 1.4); paravertebral tubercles between rear of head and cloaca 43—56 (49.6 ± 3.3), between axilla and groin 25—32 (29.1 ± 2.2); scales around midbody 92—116 (103 ± 7.4); ventral scales from gular region to cloaca 67—87 (73.5 ± 5.7), across midbody 26—36 (29.4 ± 3).

In life ( Fig. 6 ), dorsal ground color of head, body and limbs of paratypes varies between yellowish-white, grayish-white, cream and pale brown; irregular median or dark brown flecks and blotches on the dorsal surface of the body and limbs may either be almost lacking or quite prominent and forming transverse dorsal bands or a more or less reticulate pattern, that may be discontinuous in some specimens resulting in a pale cream or light brown middorsal line; postnasal and postorbital brown stripe lacking in most specimens; ventral surface of head and body cream or yellowish, with or without brown mottling; dorsal surface of unregenerated tails with alternating cream or light brown, median brown and dark or blackish brown blotches; ventral surface of tail varying from uniformly cream or yellowish to a grayish brown reticulation with strong brown mottling; pupil silver-gray or slightly bronzecolored.

In ethanol, dorsal ground color of head, body and limbs of paratypes varies between pale brown and cream; ventral surface of head and body cream; ventral surface of tail varying from uniformly cream to grayish with strong grayish-brown mottling.

Etymology. The specific epithet pachamama is derived from the Quechuan language and can be translated as Mother Earth. It represents the name of a goddess revered by indigenous people from South America, and in particular from Andean regions. According to many believers, problems will arise when people take too much from nature, because they are stealing it from Pachamama . This name was chosen to emphasize the need for protected areas along the Marañón river and blames the numerous ongoing mining and dam construction activities for exploiting this unique habitat.

Distribution and natural history. Phyllodactylus pachamama sp. nov. is endemic to the upper valley of the Marañón River and some of its tributaries in the Northern Peruvian Andes from Zapatalgo, Province of Utcubamba, Department of Amazonas to San Vicente, Province of Bolívar, Department of La Libertad. It inhabits the seasonally dry forest, occurring in xeric habitats dominated by less dense shrubs and cacti (e.g. Armatocereus, Browningia, Espostoa ) at elevations between 521—1432 m a.s.l.

Individuals of this nocturnal gecko were active between 6.25 pm and 11.35 pm and mostly found on rock walls or house walls in heights up to about 3 m above the ground. Some individuals were also found running over sandy ground next to the Marañón River. Only one male individual (CORBIDI 5703) was found during the day hidden under some logs. During their active hours, air temperatures ranged from 23.0 to 29.7° C, substrate temperatures ranged from 20.7 to 30.0° C and air humidity ranged from 41 to 71%.

Aurich et al. (2011) examined the stomach contents of 8 individuals of P. pachamama from Balsas (referred to as P. reissii ), which contained exclusively arthropod material, dominated by coleopterans, which made up 40% of all prey items.

Three gravid females (CORBIDI 5700; ZFMK 90885, 91725) were collected in mid April (2009 and 2010), each contained two eggs. On 10 July 2005, a gravid female (ZFMK 85005) was collected which contained only one egg. Two juveniles with a SVL of 31 mm (ZFMK 90889) and 32 mm (ZFMK 90891) were found in December 2009. Most gravid females of the species examined by Aurich et al. (2015) also contained two eggs.

In Balsas and San Vicente this species occurs sympatrically with the congeners P. delsolari and P. thompsoni , and with Phyllopezus maranjonensis , whereas in Zapatalgo it was found sympatrically with the sphaerodactylid Pseudogonatodes barbouri .

Figures

FIGURE 5. Dorsal and ventral views of the holotype of Phyllodactylus pachamama sp. nov. (ZFMK 90886). Scale bars represent 5 mm.

FIGURE 6. Phyllodactylus pachamama sp. nov. from the type locality (Balsas, Amazonas, Peru) in life.

Tables

TABLE 3. Morphological variation of the characters examined in Phyllodactylus reissii and related species. Values shown are mean value, standard deviation, and range. Characters marked with an asterisk are metric (in mm). See Materials and Methods for a description of abbreviations for morphological characters.

	P. interandinus	P. johnwrighti	P. magister	P. pachamama sp. nov. P. reissii
	n = 40	n = 16	n = 49	n = 22 n = 39
SVL*	39.88 ± 4.77 25.0–49.0	30.64 ± 5.95 19.0–38.5	51.68 ± 9.48 31.0–69.0	60.77 ± 13.52 51.10 ± 11.52 31.0–77.0 23.6–73.4
HL*	12.14 ± 1.61 7.4–17.3	10.25 ± 1.45 7.4–12.3	15.84 ± 2.70 10.0–20.3	18.53 ± 3.76 12.81 ± 3.45 10.0–23.0 5.5–18.5
HW*	6.30 ± 0.74 4.2–7.4	5.16 ± 0.79 3.4–6.2	8.51 ± 1.49 5.0–10.9	10.67 ± 2.49 9.3 ± 2.39 5.5–17.0 5.0–14.5
EYE*	2.35 ± 0.40 1.3–3.3	2.02 ± 0.33 1.4–2.5	3.00 ± 0.60 1.3–4.1	3.59 ± 0.78 3.15 ± 0.74 2.0–4.9 1.6–4.6
EAR*	0.83 ± 0.19 0.5–1.2	0.73 ± 0.23 0.2–1.2	1.46 ± 0.47 0.6–2.7	1.81 ± 0.48 1.12 ± 0.47 0.9–2.6 0.3–2.7
DEE*	3.57 ± 0.40 2.6–4.3	2.91 ± 0.69 1.5–4.0	4.90 ± 0.89 3.2–6.4	5.65 ± 1.32 4.45 ± 0.93 2.9–7.5 2.8–6.1
AGL*	17.39 ± 2.84 9.5–22.0	12.43 ± 3.38 6.6–18.4	21.27 ± 4.65 12.0–31.0	25.19 ± 6.02 21.80 ± 5.32 12.0–35.0 10.1–33.5
sl	7.87 ± 0.73 7–10	8.75 ± 1.34 6–11	9.98 ± 1.44 7–13	10.18 ± 1.40 8.44 ± 1.33 8–13 7–12
il	6.54 ± 0.64 5–8	7.19 ± 0.54 6–8	8.20 ± 1.24 6–12	8.73 ± 1.03 7.41 ± 0.75 7–11 6–9
lff	10.18 ± 1.06 8–13	9.40 ± 1.35 8–12	12.39 ± 1.00 10–14	12.50 ± 0.86 11.18 ± 0.85 10–14 10–13
lft	12.63 ± 1.03 11–16	10.53 ± 1.92 7–14	14.73 ± 1.11 12–18	15.09 ± 0.81 13.26 ± 0.99 14–17 12–16
dom	77.05 ± 6.50 57–88	73.31 ± 6.50 65–93	90.33 ± 7.53 78–114	103.00 ± 7.42 84.13 ± 7.56 92–116 69–102
in	0.10 ± 0.30 0–1	1.06 ± 0.85 0–3	2.06 ± 0.56 0–3	2.09 ± 0.29 1.33 ± 1.11 2–3 0–3
tdt	16.38 ± 2.14 12–22	13.38 ± 1.20 10–14	14.20 ± 1.27 12–16	13.18 ± 1.37 12.33 ± 1.03 10–16 10–14
ldt	67.85 ± 6.47 54–83	45.06 ± 2.54 40–50	49.16 ± 3.74 40–60	49.64 ± 3.30 45.62 ± 5.27 43–56 31–57