Phyllodactylus magister

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 : 53-55

Phyllodactylus magister

Phyllodactylus magister Noble, Occasional Papers of the Boston Society of Natural History 5:110. ―1924

Phyllodactylus magister (partim) — Burt & Burt, Bulletin American Museum of Natural History, 61: 250. — 1931

Phyllodactylus reissii (partim) — Dixon & Huey, Los Angeles County Museum Contributions in Science 192: 50. ―1970

Phyllodactylus reissii (partim) — Peters & Donoso-Barros, United States Nat. Mus. Bull. 297: 242. — 1970

Phyllodactylus reissii (partim) — Peters & Donoso-Barros, Smithsonian Institution Press, Washington D.C. & London: 242. — 1986

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. MCZ 17974, adult male, collected in deserted huts in the Chinchipe valley near Perico, Province of Jaén, Department of Cajamarca by G.K. Noble during the Harvard Peruvian Expedition in September 1916. (Pictures of the preserved holotype are available at http://mczbase.mcz.harvard.edu/MediaSet.cfm?media_id=10875)

Paratypes (153). All paratypes collected by G.K. Noble during the Harvard Peruvian Expedition in 1916 form the Province of Jaén , Department of Cajamarca: MCZ 18126—18129, 18141 , 18142, 126238—126375, 169013 from Bellavista ; MCZ 18143—18150 from Perico.

New collected material (47). See Appendix 2.

Diagnosis and comparison. According to our data Phyllodactylus magister reaches a maximum SVL of 69 mm. Compared to other species of mainland South America it is thus slightly smaller than P. pachamama , P. delsolari , P. dixoni , P. reissii and P. ve n t r a l i s (all exceeding 70 mm SVL) and larger than P. angustidigitus (SVL ≤ 57 mm), P. clinatus (SVL ≤ 46 mm), P. gerrhopygus (SVL ≤ 56 mm), P. heterurus (SVL of the single known specimen is 38 mm), P. inaequalis (SVL ≤ 42 mm), P. interandinus (SVL ≤ 49 mm), P. johnwrighti (SVL ≤ 44 mm), P. kofordi (SVL ≤ 46 mm), P. leoni (SVL ≤ 55 mm), P. lepidopygus (SVL ≤ 55 mm), P. microphyllus (SVL ≤ 58 mm), P. pumilus (SVL ≤ 51 mm), P. sentosus (SVL ≤ 56 mm), and P. thompsoni (SVL ≤ 42 mm). The species can be distinguished from P. thompsoni by the lack of an enlarged postanal plate. It further differs from P. angustidigitus , P. gerrhopygus , and P. heterurus by the absence of a preanal plate. By having 12–16 well-defined rows of enlarged, trihedral keeled tubercles, P. m a gi s t e r 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). Having 40—60 paravertebral tubercles between rear of head and posterior edge of thigh differentiates the species from P. ko f o rd i (31—36), P. interandinus (54—83) and P. sentosus (26—31). Phyllodactylus magister can further be differentiated from P. angustidigitus , P. clinatus , P. gerrhopygus , P. inaequalis , P. lepidopygus , and P. microphyllus by the presence of tubercles on the tibia. By the absence of tubercles on the forearm it can be differentiated from P. dixoni , P. kofordi , P. sentosus , and P. ventralis and by the absence of tubercles on the tail it further differs from P. heterurus , P. kofordi , P. pumilus and P. sentosus . In contrast to P. angustidigitus , P. microphyllus , and P. sentosus the species possesses large terminal lamellae. By having 12— 18 (mean 14.7) lamellae under the fourth toe it can be distinguished from P. inaequalis (10—12, mean 10.7), and P. leoni (9—13, mean 10.9). It further differs from P. reissii and P. pachamama by the presence of small tubercles on the thigh in about half of the specimens (always absent in P. reissii and P. pachamama ), and besides it differs from P. pachamama by having always 2 postmentals (2—4, mean 2.5 in P. pachamama ), and by an on average smaller number of scales around midbody (78—114, mean 90.3 in P. m ag i s t e r vs. 92—116, mean 103.0 in P. pachamama ). Compared to P. reissii the anterior edge of the ear opening is smooth or only slightly denticulated in most specimens (strongly denticulated with pointed scales in P. reissii ).

Redescription. The following redescription is based on the adult female ZFMK 90883 from the type locality Perico. Head distinct from neck, covered with small granules that are largest on snout and smallest on interparietal region; rostral 2.8 mm wide, 1.5 mm high, rectangular, with a median groove along its posterior one third; 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; granules in loreal region about three times larger than scales in interorbital region; 21 scales across snout at level of third supralabial, 17 at level of second supralabial; 11 supralabials, six to a point below the center of eye; eyes large, pupil vertically elliptical, with corrugated edges; orbital diameter 3.3 mm; eyelid with three visible rows of granules on both sides, granules of median row strongly pointed in posterior part; 23 midorbital granules; interorbital distance 2.0 mm; 22 granules between posterior edge of eye and anterior edge of ear, eye to ear distance 5.0 mm; ear opening elliptical, posterodorsally oriented, 1.9 mm high and 0.7 mm wide, its anterior edge almost smooth and the posterior edge only very slightly denticulated; rear of head intermixed with small, round, pulvinate tubercles; mental bell-shaped, larger than infralabials, about as long as wide; two postmentals, in contact with first pair of infralabials laterally and seven scales posteriorly; nine infralabials, six to a point below center of eye, first two pairs largest, adjacent pairs of infralabials continuously 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 14 longitudinal rows of enlarged, trihedral and slightly keeled tubercles at midbody; eight rows of tubercles reach to occiput and six rows reach to base of tail; paravertebral row with 45 tubercles between occiput and cloaca, 29 between axilla and groin; paravertebral rows separated from each other by 4—7 granules; tubercles in paravertebral row usually separated from each other by 1—2 granules; 1—4, mostly 2 granules between other longitudinal tubercular rows; 90 scales around the midbody; ventral surface with smooth, imbricate scales, constantly increasing in size towards vent, 65 from posterior gular region to cloaca, 28 trǝnsversely 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, with at least small tubercles; dorsal surface of tibia scattered with small cone-shaped or pulvinate tubercles, which are much smaller than the large tubercles on the back and about 3—4 times larger than adjacent granules on tibia; subdigital lamellae on left fingers (I—V) 8—10—11—12—9; subdigital lamellae on left toes (I—V) 9—11—13—14—13; terminal and penultimate lamellae paired, terminal lamellae greatly enlarged, truncated, the tip of the claw hardly visible, slightly projecting terminal pads; tail complete and unregenerate, 1.2 times the SVL, covered with smooth and imbricate scales, heterogeneous in size; most scales in median row of subcaudals slightly to moderately enlarged.

Measurements in mm. — Snout-vent length 53; tail length 63.7; axilla-groin length 26; right forelimb length 21.3; right hindlimb length 24; head length 15.5; maximum head width 10.8; midorbital head width 8.8; head height 6.3.

Coloration of ZFMK 90883 in ethanol. Dorsal ground color of head, body, limbs and tail pale grayish brown or cream, most scales mottled with dark brown, forming some irregular small brown flecks and blotches on the dorsal surface of the body and limbs; no facial stripe visible; ventral surface of head and body cream, with a trace of a brown mottling on about half of the scales; subcaudal ground color cream, with grayish brown mottling that becomes denser towards tip; lamellae under fingers and toes pale grayish.

Variation (see also Table 3 ; based on the paratypes MCZ 18142 and MCZ 18145 and 47 newly collected specimens). — SVL of adult males ranges from 51–69 mm (58.6 ± 4.2, n = 16), and SVL of adult females ranges from 50–66 mm (54.4 ± 4.2, n = 9). Tail length/total length is 0.45–0.56 (0.52 ± 0.02, n = 36). Male specimens possess 3—5 (3.6 ± 0.69) postanal tubercles on each side. The number of postmentals is 2 in all specimens. In the majority of specimens postmentals contacting first infralabial on each side, scales bordering postmentals 5—7 (6.0 ± 0.62). Internasals 0—3 (2.1 ± 0.56). Lamallae on fourth finger 10—14 (12.4 ± 1.00); lamellae on fourth toe 12—18 (14.7 ± 1.11); supralabials 7—13 (10.0 ± 1.44), to a point below the center of eye 6—8 (7.2 ± 0.73); infralabials 6—12 (8.2 ± 1.2), to a point below the center of eye 6—8 (6.2 ± 0.53); loreals between postnasal and anterior edge of orbit 9—15 (11.8 ± 1.45); granules between posterior edge of eye and anterior edge of ear 21—30 (25.1 ± 2.55); anterior edge of the ear opening is smooth or only slightly denticulated in most specimens; scales across snout at level of third supralabial 20—27 (22.6 ± 1.62), at level of second supralabial 14—18 (15.8 ± 1.14); midorbital granules 19—26 (22.7 ± 1.76); longitudinal rows of dorsal tubercles 12—16 (14.2 ± 1.27); paravertebral tubercles between rear of head and cloaca 40—60 (49.2 ± 3.74), between axilla and groin 24—35 (28.4 ± 2.94); tubercles on tibia are present in all specimens, but may be very small and very few in some specimens; very small and few tubercles on thigh present in about half of the specimens; scales around midbody 78—114 (90.3 ± 7.53); ventral scales from gular region to cloaca 63—80 (70.0 ± 4.21), across midbody 23—32 (26.8 ± 1.77).

In life, dorsal ground color of head, body and limbs varies between grayish-white, yellowish-cream, cream and pale brown; irregular whitish, brown or dark brown flecks and blotches on dorsal surface of the body and limbs may either be hardly visible or quite prominent and forming transverse dorsal bands or a more or less reticulated pattern (in the majority of specimens), that may be discontinuous in some specimens resulting in a pale cream or light brown middorsal line; dark brown line extending laterally on each side from nostril through eye to above insertion of the forearm conspicuous in most specimens, faint or absent in others; ventral surface of head and body cream to bright yellow, 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 bright yellow to a grayish brown with strong brown mottling; pupil golden brown or coppery; tails of most juveniles annulated with yellow and brown.

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

Distribution and natural history. Phyllodactylus magister is endemic to the seasonally dry forest vegetation of the upper Marañón River basin and some of its tributaries in the Northern Peruvian Andes in the Departments Cajamarca and Amazonas from Perico to Puerto Malleta, at elevations between 419—1247 m a.s.l..

The habitat of Phyllodactylus magister is composed of drought-resistant trees (e.g. Acacia, Anadenanthera, Ceiba, Cordia, Prosopis ), dense shrubs (e.g. Croton, Mimosa ) and ground vegetation layer (e.g. Opuntia , Poaceae ).

During daytime surveys we found several specimens of this nocturnal species hidden under stones or inside deserted huts. Active individuals were recorded between 6.45 pm and 0 1.00 am, primarily on rock walls, sandy walls besides paths, roads or dry creek beds, as well as house walls from ground level up to about 2.5 m above the ground. Air temperatures during the active hours of this species ranged from 19.2 to 29.9° C, substrate temperatures ranged from 17.4 to 31.9° C and air humidity ranged from 52 to 75%.

Gravid females of P. m ag i s t e r were collected end of May (ZFMK 88739), and in mid December (CORBIDI 5708, ZFMK 90898), containing two eggs, except for ZFMK 90898 which had only one egg. On the 8th of May 2008 at 11.15 pm a gravid female was discovered (not collected) directly after it had deposited an egg in the soft soil of the sandy walls of a dry creekbed. One juvenile with a SVL of 31 mm (ZFMK 90881) was found in March 2009 and another juvenile with a SVL of 33 mm (CORBIDI 1812) was found in May 2008. Subadults with SVL between 37—45 mm were collected in March 2009 (ZFMK 90880, 90882), April 2009 (CORBIDI 5696, 5697), May 2008 (CORBIDI 1829, 1833, 1836), July 2008 (CORBIDI 1819, ZFMK 88741), and in December 2009 (ZFMK 90895, 90896, 90901, 90902).

We found Phyllodactylus magister in sympatry with the congener P. interandinus in Jaén, Bellavista and Puerto Malleta , and according to Dixon & Huey (1970) the two species also occur sympatrically in Bagua Chica, Bagua Grande, and Perico. In Jaén and Santa Rosa de la Yunga, we observed P. magister in the same habitat as the sphaerodactylid species Gonatodes atricucullaris , and Noble (1921) also adds Bellavista as a contact zone of both species. According to him, a second sphaerodactylid species, Pseudogonatodes barbouri , also occurs in Bellavista, but we did not find this species in sympatry with P. m ag i s t e r in Perico.