Paragehyra austini, Crottini & Harris & Miralles & Glaw & Jenkins & Randrianantoandro & Bauer & Vences, 2015

Crottini, Angelica, Harris, D. James, Miralles, Aurélien, Glaw, Frank, Jenkins, Richard K. B., Randrianantoandro, J. Christian, Bauer, Aaron M. & Vences, Miguel, 2015, Morphology and molecules reveal two new species of the poorly studied gecko genus Paragehyra (Squamata: Gekkonidae) from Madagascar, Organisms Diversity & Evolution 15 (1), pp. 175-198 : 190-193

Paragehyra austini sp. nov.

Refer to Figs. 1d , 2 , 4b and 6 .

Holotype ZSM 339/2005 (FGZC 2366), adult male with everted hemipenes and open ventral portion of the right forelimb, collected at Grotte Ampasy ( Andohahela National Park ), 24° 32.663′ S; 46° 41.249′ E; 796 m a.s.l., western slope of the Andohahela massif, Tolagnaro ( Fort Dauphin ) Fivondronana , Toliara province, southeastern Madagascar, on 26th January 2005 by Parfait Bora, Frank Glaw and Miguel Vences.

Paratypes ZSM 338/2005 (FGZC 2355), young female or unsexed subadult in good condition and with open ventral portion of the right hindlimb; ZSM 340/2005 (FGZC 2367), adult male with everted hemipenis and with autotomised tail (tail preserved with the specimen); UADBA uncatalogued (FGZC 2365), juvenile; UADBA uncatalogued (FGZC 2368), adult male; all collected at the same locality and dates as the holotype.

Diagnosis A Paragehyra with no enlarged tubercles on dorsolateral surfaces of the body (character BT); no tubercles on dorsal surfaces of limbs (character TDL); and no tubercles on dorsolateral surface of tail (character TT); enlarged infralabials diminish gradually in size posteriorly (character I); small lateral chin scales extend anteriorly along each side, excluded from contact with first infralabials and first postmentals, in contact with second infralabials (character C); ventral scales on distal (tibial) segment of hindlimb slightly larger than the other ventral scales (character VET); scales on preaxial-ventral portion of pes between end of tibia and base of digit I relatively small, seven scales visible (character SPP); three to four small, subdigital scales between enlarged basal scales and terminal distal pad on digit I of manus and pes (character SS); claw-bearing segment of digits II–V of manus and pes with three to seven subdigital scales (character SSC); smooth dorsal scales equal in size than ventrals (character DO); smooth pigmented ventral scales (character VE); smooth pigmented subcaudal scales (character SC); triangle-shaped mental scale (character ME); first large

P. petiti ZSM 592/2000 with bell-shaped mental scale and the Paragehyra sp. aff. petiti 1 from Tsingy de Bemaraha (UADBA 28056) showing the modified bell-shaped mental scale postmental scales in contact for more than the 50 % of their length (character 1 PM).

P. austini sp. nov. is most similar to P. gabriellae from which it is differentiated in five characters: BT (absence of enlarged tubercles dorsolaterally on the body vs. presence), VET (ventral scales on distal (tibial) segment of hindlimb slightly larger than the other ventrals vs. ventral scales on distal (tibial) segment of hindlimb of normal size), SS (3–4 vs. 7–10 smaller subdigital scales between enlarged basal scales and distal pad on digit I of manus and pes), SSC (subdigital scales on claw-bearing segment of digit II–V of manus and pes with distalmost scale markedly larger than the others vs. numerous small scales increasing gradually in size distally), 1PM (first large postmental scales in contact for more than the 50 % of their length vs. first large postmental scales in contact for equal or less than the 50 % of their length).

P. austini sp. nov. differs from P. petiti in 12 characters: BT (absence of enlarged tubercles dorsolaterally on body vs. 10 longitudinal rows of enlarged tubercles dorsolaterally on body), TDL (absence vs. presence of enlarged tubercles on distal segment of forelimbs), TT (absence vs. presence of transverse rows of enlarged tubercles on tail), I (enlarged infralabials diminish gradually in size posteriorly vs. first four to six infralabials much larger than remainder), VET (ventral scales on distal (tibial) segment of hindlimb slightly larger than the other ventrals vs. five to six enlarged ventral scales on distal (tibial) segment of hindlimb), SPP (seven small scales along preaxial-ventral border of pes vs. five large scales along preaxial-ventral border of pes), SS (three to four small subdigital scales between enlarged basal scales and terminal distal pad on digit I of manus and pes vs. two to three small scales), SSC (subdigital scales on claw-bearing segment of digit II–V of manus and pes with distalmost scale markedly larger than the others vs. large and almost equally in size scales), DO (smooth dorsal scales equal size to ventrals vs. smooth dorsal scales smaller than ventrals), VE (smooth pigmented ventral scales vs. smooth unpigmented ventral scales), SC (smooth pigmented subcaudal scales vs. smooth unpigmented subcaudal scales), ME (triangle-shaped mental scale vs. bell-shaped mental scale).

P. austini sp. nov. differs from P. felicitae sp. nov. in eight characters: BT (absence of enlarged tubercles vs. 12 longitudinal rows of enlarged tubercles dorsolaterally on body), TDL (absence of enlarged tubercles on dorsal surfaces of limbs vs. presence), TT (absence of transverse rows of enlarged tubercles on tail vs. presence), VET (ventral scales on distal (tibial) segment of hindlimb slightly larger vs. 6 enlarged ventral scales on distal (tibial) segment of hindlimb), SPP (7 small scales along preaxial-ventral border of pes vs. 6 large scales along preaxial-ventral border of pes), SS (3–4 small, subdigital scales between enlarged basal scales and terminal distal pad on digit I of manus and pes vs. 2–3 small scales), SSC (subdigital scales on claw-bearing segment of digit II–V of manus and pes markedly larger than the others vs. scales almost subequal), VE (smooth pigmented ventral scales vs. smooth unpigmented ventral scales).

Remarks In addition to the morphological characters, P. austini differs from all other species by substantial genetic differentiation in mitochondrial and nuclear markers.

Description of the holotype Refer to Figs. 1d and 2 ; measurements are in Tables 2 and 3. Male in good condition with fully everted hemipenes and open ventral portion of the right forelimb. Two scratches on the dorsal body. Body and head flattened dorsoventrally. Head slightly narrower than the widest part of the body (cf. HWand BW), around 1.5 times longer than wide (cf. HL and HW in Table 1). Snout long and with a rounded tip. Pupil round. Eye large (ED), diameter 0.39 times snout length (SnL). Ear opening (EO) elliptical, with nearly vertical long axis, slanting slightly posteriorly in the ventrodorsal direction, smaller than eye, vertical diameter 0.56 times eye diameter. Body (snout-vent length [SVL] minus head length [HL]) relatively short, around 2.2 times longer than head. Limbs and digits relatively long, forelimb (FL) when extended forward reaches midway between anterior edge of eye and nostril, when extended posteriorly, reaches three quarters of distance to groin, hindlimb (HiL) reaches anterior axilla. Tail regenerated (TAL) after the fourth whorl, 1.14 times as long as SVL, subcylindrical, flattened dorsoventrally, especially at the base, and less so distally, and tapering to a sharp point. Quadrangular rostral scale as wide as mental scale with a short, incomplete dorsal vertical groove extending downward for around one fourth the distance from dorsal edge to lip. Concave groove between nasals, one large internasal, nasals not in contact (IN). Nostrils in contact with rostral, nasals, three postnasals and first supralabials. Number of supralabials (SL) (right/left) 8/9; number of infralabials (right/left) 8/8. Supralabials and infralabials decreasing gradually in size posteriorly.

Mental (ME) large, triangular in contact with first infralabials and bordered posteriorly by a pair of large elongate irregular hexagonal to pentagonal postmentals. A row of six smaller enlarged scales (here called second postmentals [2PM]) separates the mental and postmentals from the smaller chin and throat scales. Large postmentals (1PM) (longer than mentals) in contact for more than the 50 % of their length and in contact with mental and first infralabials anteriorly, with each other posteriorly, with a pair of enlarged second postmentals (2PM) posterolaterally, and with two enlarged second postmentals (2PM) posteriorly. Slightly enlarged chin scales present between second postmental and infralabials on each side; chin scales decrease gradually in size posteriorly. Lateral chin scales extend forward along each side, right anteriormost chin scale in contact with right second postmentals and right second infralabials; left anteriormost chin scale in contact with left second postmentals and left second infralabials.

Dorsal scales (DO) consist mostly of small smooth granular scales equal in size to ventral scales. Uniform scalation on all dorsal surface. Enlarged tubercles absent from all dorsal surfaces (BT, TDL, TT). Regenerated tail with four discernible whorls. Tail base wide (TBW). Distal to base, dorsal scales of tail small and cycloid, becoming slightly imbricate posteriorly. Ventral scales (VE) smooth, cycloid, varying in size and degree of overlap. Generally non-imbricate in regions where scales are relatively small and imbricate where scales are larger. Throat scales small, circular and juxtaposed. Scales just ventral to posterior infralabials enlarged. Throat covered by small granular scales. Imbricate, cycloid scales begin behind throat and cover chest, belly, hindlimb and tail. Ventral surface of forelimbs covered with granular scales on both proximal and distal segments. Ventral surface of pelvis and thigh (proximal portion of hindlimbs) covered with imbricate cycloid scales. Ventral, distal segment of hindlimbs (VET) with slightly larger and numerous, imbricate, cycloid scales. Subcaudal scalation as follows: tail base with imbricate and cycloid scales, smaller than the following subcaudals. First whorl with one set of enlarged cycloid scales proximally and two wide plates distally; whorls two and three with a row of three, wide, undivided plates; whorl 4 with a row of two, wide, undivided plates; regenerated portion with a row of wide, undivided plates.

Dorsal scales on tail larger than the dorsal scales on body, ventral plates separated ventrolaterally by one to three longitudinal rows of large, imbricate, cycloid scales. Scales immediately adjacent to cloacal opening much smaller than surrounding scales.

Preaxial border of palm and digit I of manus covered with numerous small scales. Seven relatively small, imbricate, cycloid scales along preaxial-ventral border of pes on each side (SPP). Granular or weakly imbricate scales cover palms and bases of digits II–Von manus and pes. Pads of digits II–V confined to distal two thirds of pad-bearing segment of digits. Pads bearing rows of undivided scales. Numbers of (transversely enlarged) subdigital lamellae in order on digits II–V (SLMP): left manus 8,8,8,8; left pes 8,8,9,8. Digit I of manus and pes has three kinds of subdigital scales linearly arranged along digit: a basal row of enlarged scales, an intermediate row of small paired scales (SS) and a large terminal quadrangular pad. Numbers of these, in order, are as follows: left manus, 2,3,1; left pes 3,3,1. Terminal pad of digit I pilose and lies before the claw. Claw-bearing segment of digits II–V has a ventral row of small, imbricate, cycloid, subdigital scales from base of digit to base of claw (SSC): distalmost scale markedly larger than the others: numbers of these, in order, are as follows: left manus 4,3,3,5; left pes 4,4,5,6. Comparative finger and toe length 1<2<5<3<4. Two precloacal pores (PCP), one left and one right. Pores separated by 1 poreless scale.

Testes white, enlarged; left testis 6 mm long and 3.2 mm wide. Vas deferentia enlarged ( Fig. 5d, e ). The hemipenes show a bilobed structure with clear differentiation between truncus and apex. The sulcus spermaticus is long and not very deep. Spines and/or denticulated papillae entirely absent. Truncus slender and large sulcated apex divided in two welldefined smooth lobes ( Fig. 5d, e ).

Colouration Colour after 9 years in alcohol similar to that in life. Ground colour of head, body, tail, hands, feet and dorsal parts of limbs dorsally and laterally light grey-brownish with darker brownish blotches and bands on dorsal surface of body and limbs. A longitudinal row of nine midline dorsolateral brownish blotches on dorsal surface of body with the 3rd−7th with a lighter central spot along midline. A grey and brownish regular pattern along the dorsolateral portion of the tail. A black band from anterior border of eye continuing frontward until one third to nostrils. Chin, margin of lower jaws, lower cheeks, lateral throat regions, palms and subdigital areas light brownish ( Figs. 2 and 4b). Ventral part of body with uniformly white-brownish scales along the midportion of chest and belly and with light brownish scales ventrolaterally; light brownish scales along the ventral portion of the tail. Supralabials and infralabials light brownish with numerous scattered melanophores.

Variation Based on ZSM paratypes, for measurements, see Tables 2 and 3. Colouration of ZSM 340/2005 overall similar to that of the holotype but with a faded brownish colour and a colour pattern less discernible. Colour and pattern of ZSM 338/2005 slightly different from the holotype.

In ZSM 340/2005, the supralabials on the right side are 10, three precloacal pores are visible (one on the right side and two on the left side), nine infralabials on the left side and four subdigital scales between enlarged basal scales and distal pad on digit I of manus.

In ZSM 338/2005, nine supralabials on the right, two precloacal pores on the right side of the body, nine infralabials on the right side are four subdigital scales between enlarged basal scales and distal pad on digit I of manus.

For the variation in the number of subdigital lamellae under pad-bearing portion of digits II–V of manus and pes, refer to Table 4 (character SLMP); for the variation in the number of subdigital scales in claw-bearing segment of digits II–V of manus and pes, refer to Table 5 (character SSC).

Regenerated tails differ markedly in colour compared to original tails. The banded pattern of original tails is replaced with longitudinal streaks of brown and white on regenerated tails.

Everted hemipenes are available for one additional male (ZSM 0340/2005; Fig. 5f ). Hemipenes of the two males are generally similar in structure.

Distribution, conservation and proposed IUCN Red List status The new species is currently known only from the type locality on the western slopes of the Andohahela massif which is included in the Andohahela National Park. Further investigations in the surrounding areas are required to define the actual distribution of P. austini sp. nov. Due to the very limited knowledge on this species, we suggest that its conservation status should be “Data Deficient” under IUCN criteria ( IUCN 2001).

Habitat and habits The five type specimens of P. austini sp. nov. were found during the day on the walls of a cave that was used for a short break when climbing up from Esomony to the Andohahela reserve . The species was abundant in the cave, but no further field notes were taken. The climate and vegetation at the type locality were in a narrow transitional band between arid low altitude thornbush savannah (Esomony) and humid high altitude heathland mixed with montane rainforest (Andohahela massif) .

Etymology The name “ austini ” is a personal noun in the genitive case. D. J. Harris dedicates this new species in honour of Mr. Austin O’ Malley.

Figures

Fig. 1 a Paragehyra petiti from near Toliara; b Paragehyra felicitae sp. nov., male holotype (ZSM 1611/2010) from Anja reserve; c Paragehyra gabriellae from eastern slopes of Andohahela; d Paragehyra austini sp. nov., male holotype (ZSM 339/2005) from Grotte Ampasy (near Esomony), showing colouration in life

Fig. 2 Paragehyra austini sp. nov., male holotype (ZSM 339/ 2005) (above) in dorsal and ventral view. In the dorsal view, the absence of enlarged tubercles along the body and limbs and on the tail (characters BT, TDL, and TT) is evident; while in the ventral view, the uniform smooth surface characterised by pigmented scales along the body, limbs and tail (characters VE and SC) is visible. Paragehyra felicitae sp. nov., male holotype (ZSM 1611/2010) (below) in dorsal and ventral view. In the dorsal view, it is possible to observe the 12 longitudinal rows of enlarged tubercles (character BT), and the presence of enlarged tubercles on the distal and proximal segments of the limbs (TDL) and on the dorsal surface of tail (character TT). In the ven- tral view, the uniform smooth surface characterised by unpigmented scales along the body and the limbs (character VE) and by pigmented scales along the ventral surface of tail (character SC) is shown

Fig. 6 Mental region showing differences in scalation in Paragehyra felicitae sp. nov. ZSM 1611/2010 (HT) with triangle-shaped mental scale, P. austini sp. nov. ZSM 0339/2005 (HT) with triangle-shaped mental scale, P. gabriellae ZSM 0114/2004 with triangle-shaped mental scale,

Fig. 5 a Right hemipenis of P. felicitae ZSM 1611/2010 (HT); b left hemipenis of P. felicitae ZSM 1612/2010 (PT); c right hemipenis of P. felicitae ZSM 1610/2010 (PT); d left hemipenis of P. austini ZSM 0339/2005 (HT); e right hemipenis of P. austini ZSM 0339/2005 (HT); f right hemipenis of P. austini ZSM 0340/2005 (PT). Bars 2 mm