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Lepidodactylus pantai, new species (Figures 2,3,4) Holotype. (Museum Zoologicum Bogoriense.Lace.14062, Field number ALS 534) An adult male collected by ALS, BRK, and UA from beachside rocks at Pasir Panjan Beach, Desa Ohoililir, Kei Kecil, Maluku, Indonesia a few hours after sunset at 5.646671° S, 132.638312° E (WGS84) on 16 October, 2011. Liver tissue is preserved in duplicate in RNA-Later at Museum Zoologicum Bogoriense (MZB.Lace.14062) and Museum of Vertebrate Zoology (MVZ 273691). Paratypes. A series of ten additional specimens were collected at the same locality and time as the holotype (MVZ tissue number and field number in parenthesis): MZB.Lace.14064 (MVZ 273686; ALS 502), MZB.Lace.14065 (MVZ 273687; ALS 505), MZB.Lace.14066 (MVZ 273688; ALS 530), MZB.Lace.14067 (MVZ 273689; ALS 531), MZB.Lace.14068 (MVZ 273690; ALS 533), MVZ 273692 (ALS 501), MVZ 273693 (ALS 503), MVZ 273694 (ALS 504), MVZ 273695 (ALS 532), MVZ 273696 (ALS 535). Diagnosis. A moderate-sized bisexual species of Lepidodactylus, SVL 36.9–40.5 (mean = 38.3) mm for five adult males and 32.0–40.5 (mean = 37.4) for five adult females, distinguished from other species by the following combination of characters: 113 rows of scales around the midbody; subdigital scansors 10–12 on toe IV, and 7–9 on toe I; terminal scansor is divided on digits II through V on both the fingers and toes; 3 scansors on 4th toe divided or deeply notched; interdigital webbing small, less than 1/5th the 4th toe length; 18–24 pores in precloacal and femoral regions of male. Tail nearly cylindrical without lateral serrations. Description of holotype. MZB.Lace.14062 (measurements in mm, after preservation). Snout–vent length 40.57; head length 11.74; head width 7.56; head height 4.48; jaw length left/right 6.06/6.34; snout–eye length 4.54; naris–eye length 3.13; naris circular, approximately 0.4×0.5; orbit diameter 2.46; eye–ear length 3.38; snout width 1.78; ear opening length×width 0.62×0.41; interorbital width 4.00; snout–forelimb length 14.13; axilla–groin distance 20.05; length of hind limb 15.13 (75.5 % of axilla–groin distance); length of forelimb 10.95; crus length 6.41; tail length 24 (entire); tail width 4.34; tail depth 3.88. Snout tapered, rounded at tip; three scales touching rostral between left and right nares; supranasals separated by three scales in contact with rostral; rostral entering nares, broader than high, 2.05×0.89 (width about 2.3 times height); no rostral cleft; nares bordered by five scales: three nasals, one rostral, and one supralabial; 34–35 interorbital scales; 11 left and 11 right supralabials, 10th below center of eye; 10 left and 10 right infralabials; mental scale distinct, triangular, its anterior width nearly equal to midline length (0.69×.0.72); Mental is bordered posteriorly by two enlarged primary postmentals, each in contact with the first infralabial. Body slightly depressed; 113 rows of scales (average of 3 counts: 109, 114, 116) around midbody, grading into granular scales on lower lateral surfaces; dorsal and lateral scales granular, without enlarged tubercles; ventral scales almost flat, hexagonal, and 2–3 times larger than dorsal scales; limbs well developed; digits moderately dilated, undersurface (Fig. 3) bearing left/right scansors as follow: fingers— I 9 /9, II 9/ 10, III 12 / 12, IV 13 / 13, V 9 / 9; toes—I 10/(not intact), II 11/ 11, III 14 / 14, IV 12 / 12, V 9 /9; distal three scansors, including the terminal one, divided on all digits except the first on fingers and toes; first digit with complete terminal and two divided subterminal scansors; all digits except first with 3 undivided or deeply notched scansors, including the terminal; all digits except first clawed; compressed claw-bearing phalanges arising from distal margin of the dilated part and extending only a short distance beyond; slight webbing 1/6th of the length of the 4th toe; digits elongate and slender, toe pads slightly enlarged. Twenty five enlarged precloacal and femoral scales; two post-cloacal spurs on each side of vent; tail entire, subcylindrical throughout length, gradually tapering to a blunt tip; lateral margins without spines or skin flanges; tail constricted at base posterior to vent; scales on tail annulate, squarish or rectangular, ventral scales about twice as large as dorsal scales; base of tail distinctly swollen by hemipenes; hemipenes everted, forked with small, flared scales on the midbody of each forked end, transitioning into even smaller distal scales about half the size. Color in preservative. Color of holotype after about three years in ethanol is similar to coloration in life (Fig. 2). Overall dorsal coloration is pale grey with some lateral streaks of darker pigment. Two pairs of dark spots are clearly visible on the dorsal surface of tail. Venter of body pale cream in color with no distinct pigmentation. Ventral surface of tail slightly darker posteriorly. Slight dark spotting on the dorsal surface of the head. A brown orbital stripe extends from the nostril to the anterior insertion of the forelimb. Three elongated spots on dorsal surface of body between the insertions of the forelimbs. Variation. The type series varies in SVL (20.6–40.5; n=11), number of precloacal pores in males (17–25; n=5), number of scansors on Toe IV (10–12, n=11), number of lamellae on Toe I (7–9, n=11), number of supralabials (11–14, n =11), number of infralabials (9–11, n=11), head width as a proportion of SVL (18–21%, n=11), and number of scale rows around the midbody (108–127, n=11). Nearly all specimens have two cloacal spurs, however two specimens have an additional cloacal spur on one side of the body. All specimens have three divided terminal scansors. Both males and females of the type series possess enlarged, pale-white, endolymphatic sacs, which are more pronounced in females. Coloration is similar among the type series (Fig. 4), however patterning is somewhat variable. Some specimens possess distinct dark chevron patterning along the dorsal surface of the body. All members of the type series have three elongate dark spots on the dorsal surface of the body between the insertions of the forelimbs. The lateral surfaces of some specimens have darker brown pigmentation, with a pale cream dorsal band running from the temporal region posterior to the base of the tail. The orbital stripe is variable in conspicuousness, though all specimens possess it to some degree. Coloration in life (Fig. 2) is very similar to that in preservative (Fig. 4), however some of the darker regions also contain reddish pigment that has been lost in preservation. As with many geckos, these individuals change their level of pigmentation with time of day and light environment. Figure 2 shows a color photograph of this species in situ (specimen uncollected) from the type locality taken on a more recent expedition in 2014. Distribution. The type series are all from Kei Kecil, however we recently collected another series of specimens from Kur Island (~ 80 km to the northwest) that appears morphologically similar, however further analysis is needed to determine the level of divergence between these two populations. Our phylogeny also indicates that specimens from Palau are remarkably similar genetically but we refrain from assigning members of that population to this species at present time. Natural history. We have experience with this species only on two islands, Kei Kecil and Kur. Specimens were associated with exposed intertidal limestone rock formations in both cases (see Fig. 1; habitat). All specimens were found at night on beachside limestone rocks (Fig. 1) or in mangroves within meters of the high tide line. Despite nearly a month of intensive nightly collecting effort (targeting geckos) on Kei Kecil we were unable to find this species in the adjacent disturbed forest. While on Kei Kecil, we resided in a small house approximately 30 meters from the high tide line and collected many other species of geckos on a small wooden shed but L. pantai sp. nov. were never encountered on this structure. This species was discovered accidentally at the end of our stay on Kei Kecil while surveying for laticaudine sea snakes in the intertidal zone. We think this shows the potential importance of sampling intertidal areas at night, something rarely done by herpetologists. All 11 specimens in the type series were collected over the span of less than one hour, and subsequent visits to Kei Kecil confirmed that this species is locally abundant, occasionally in densities of up to 1 individual per square meter. Terrestrial hermit crabs and marine isopods were also abundant on the same rocks and we believe this resource subsidy of food from the marine environment likely contribute to the locally high densities observed in this species. On Kur island, only 80 km northwest of Kei Kecil, morphologically similar geckos were also found in association with the intertidal zone and limestone rocks, but they were also numerous in a small patch of mangroves. The roots of these mangroves were submerged during high tide, but specimens were found at an even higher density on mangroves than on exposed limestone rocks (though the small mangroves were themselves growing among limestone rocks). When associated with mangroves, this species appeared to be very abundant close to the water, but trees slightly above the high-tide line were inhabited almost exclusively by another undescribed species of Lepidodactylus and members of the gekkonid genus Cyrtodactylus. Etymology. The species epithet, pantai, is the word for beach in the Indonesian national language (Bahasa Indonesia). It reflects the habitat in which the new species was discovered—a seemingly obligate association with the seashore. All specimens were found within 2 m of the high tide line. We suggest “Beach Scaly-toed Gecko” as the English common name for this species. Comparisons. The new species is the only member of the genus with divided terminal 4th toe scansors and a cylindrical tail without lateral serrations. Brown & Parker (1977) divided the genus Lepidodactylus into three groups but L. pantai sp. nov. differs from all of the previously recognized groups in having divided terminal scansors on toes 2–5 and a tail that is fully cylindrical without any fringes or compression. The presence of divided scansors distinguishes L. pantai sp. nov. from all Group I Lepidodactylus (L. listeri, L. magnus, L. manni, L. mutahi, L. oorti, L. orientalis, L. pumilus, L. browni, L. euaensis, and L. flaviocularis), which have no divided scansors (Brown & Parker 1977). The presence of divided terminal scansors on toe IV distinguishes L. pantai sp. nov. from all Group II Lepidodactylus which have undivided terminal scansors (L. gardeneri, L. guppyi, L.novaeguineae, L. pulcher, L. shebae, L. buleli, L. intermedius, L. lombocensis, L. paurolepis, L. vanatuensis, L. oligoporus, L. tepukapili, and L. ranauensis). The presence of a cylindrical tail without lateral serrations distinguishes L. pantai sp. nov. from Group III Lepidodactylus that have depressed tails with lateral serrations (L. moestus, L. lugubris, L. woodfordi, L. yami, L. aureolineatus, L. balioburius, L. christiani, L. herrei, and L. planicaudus). Lateral serrations are sometimes absent on L. yami (Ota 1987), but L. pantai sp. nov. can be further distinguished from L. yami by the number of scansors on Toe IV (10–12 in L. pantai sp. nov. versus 13–15 in L. yami) and the number of midbody scale rows (108–127 in L. pantai sp. nov. versus 145–148 in L. yami).

Limnonectes larvaepartus new species (Figs. 1, 2) Urn:lsid:zoobank.org:act: 60AA7136-89A0-4DBB-9FBC-BD0FAF8A214C This species has been referred to in the literature under the names Limnonectes larviparus [11] and Limnonectes ‘‘ovovivipar’’ [4], both of which created nomina nuda. This species also corresponds to Limnonectes sp. V in [2, 3]. Etymology The species name larvaepartus (from ‘larvae’, plural of larva, the early form of an animal, and ‘partus’, to give birth to) reflects the unique reproductive mode of this tadpole-bearing species. Holotype MZB.Amph.23755 (Field Number: BSI 0 605, see Fig. 1), an adult male, collected from Dunu Village, (0.92353o N; 122.64386o E) at 189 m elevation, Kecamatan Anggrek, Kabupaten Gorontalo, Provinsi Gorontalo, Sulawesi, Indonesia by J.A. McGuire & team, 18 October 2004. Paratypes Paratypes (n=30) are from Sulawesi Utara, Gorontalo, Sulawesi Tengah, and Sulawesi Barat Provinces: MZB 2834, a gravid female with 33 translucent tadpoles (Gosner stage 23) from the left oviduct and 2 from outside the body, from Toraut (00'33.72o N, 123'54.23o E), Bogani Nani Wartabone National Park, Kabupaten Bolaang Mongondow, Sulawesi Utara at 370 m elevation, by D. T. Iskandar, 15 August 1991; FMNH 252453, a female, from Toraut, Bogani Nani Wartabone National Park, Kabupaten Bolaang Mongondow, Sulawesi Utara by D. T. Iskandar, August 1991.MVZ 255545, 256009-11, 256013 from Desa Lombongo (‾1.43346, 120.30800), Kecamatan Suwawa, Kabupaten Bone Bolango, Bogani Nani Warta Bone National Park, Provinsi Gorontalo at 75 m elevation by J. A McGuire and team, 20 October 2004.ZRC 1.3258 (left femur removed) from Toraut, Bogani Nani Wartabone National Park, Kabupaten Bolaang Mongondow, Sulawesi Utara at 370 m elevation, by D. T. Iskandar, 15 August 1991.MZB 2835– 2841 from Toraut, Bogani Nani Wartabone National Park, Kabupaten Bolaang Mongondow, Sulawesi Utara at 370 m elevation, by D. T. Iskandar, 15 August 1991 & 12 July 1992.MVZ 255548–49 from Desa Pontak (‾2.62910, 118.99300), Kecamatan Motoling, Kabupaten Minahasa Selatan, Provinsi Sulawesi Utara at 285 m elevation by J. A. McGuire and team, 13 October 2004. MZB 3117, near Potolok river, Lolak, Bogani Nani Wartabone National Park at 350 m elevation, Kabupaten Bolaang Mongondow, Sulawesi Utara; MZB 3118, male from seashore forest near Bungbungan River, Lolak, Bogani Nani Wartabone National Park, Kabupaten Bolaang Mongondow, Sulawesi Utara.MZB 3120 from Tangkorak River, Desa Pindol, Kecamatan Lolak, Bolaang Mongondow, Sulawesi Utara, by Mumpuni, 26 June 1995; MZB 3121 from Tangaga Forest, Dudepo, Bolaang Mongondow, Sulawesi Utara, by I. Maryanto, 20 October 1995; MZB 3122 from Potolok River, Bogani Nani National Park, Lolak, Sulawesi Utara by Mumpuni, 19 June 1995; MZB 3124 from Bungbungan River, Bogani Nani National Park, Lolak, Sulawesi Utara by Mumpuni, 19 June 1995; MZB Amph.8108 from Toraut, near Bogani Nani Wartabone National Park, Sulawesi Utara.LSUMZ 84209, 84214, 84221, 84224 from Desa Torout, Bogani Nani Wartabone National Park, Kabupaten Bolaang Mongondow, Provinsi Sulawesi Utara at 267 m elevation by J. A. McGuire on 6 and 11 September 2001. doi:10.1371/journal.pone.0115884.g002 Other referred specimens LSUMZ 84218, 84219 from Desa Torout, Bogani Nani Wartabone National Park, Kabupaten Bolaang Mongondow, Provinsi Sulawesi Utara by J.A. McGuire.FMNH 130991, 106930 from Buang-buang Island, Sulawesi Utara.MZB 3119, juvenile from Seashore forest near Bungbungan River, Lolak, Bogani Nani Wartabone National Park, Kabupaten Bolaang Mongondow, Provinsi Sulawesi Utara.AMNH 167199 from Tangkoko National Park (1.570083o N, 125.156933o E), Kabupaten Minahasa, Provinsi Sulawesi Utara.MVZ 255546 from Desa Salumpaku (‾1.60757, 119.29900), Kecamatan Banawa, Kabupaten Donggala, Provinsi Sulawesi Tengah.MVZ 255547 from Desa Kelapa Dua (‾1.60757, 119.29900), Kecamatan Anreapi, Kabupaten Polewali Mandar, Provinsi Sulawesi Barat.MVZ 268426, 268428–30, 268432 from Polewali-Masawa Road (River 1) (‾2.65490, 118.93300), Kecamatan Polewali, Kabupaten Polewali Mandar, Provinsi Sulawesi Barat.MVZ 268309–10, 268313, 268317–19, 268322, 268325,MZB.Amph. 20675, 20677–80 from Desa Uaemate (Tasiu-Tibo Road; S02.61550, E119.14417), Kecamatan Kaluku, Kabupaten Mamuju, Provinsi Sulawesi Barat.MZB.Amph.20663 from Desa Kabiraan (‾2.62460, 119.14700), Kecamatan Ulumunda, Kabupaten Majene, Provinsi Sulawesi Barat. Distribution Limnonectes larvaepartus occurs across the Northern Peninsula, as well as on the western margin of Sulawesi’s Central Core (Fig. 3). We do not know the full extent of the species’ range in the Central Core because the central highlands of Sulawesi remain poorly explored herpetologically. Several genera have species range boundaries in this same general region (e.g. the flying lizards Draco spilonotus and D. walker [12], the fanged frogs Limnonectes sp. I and L. sp. D [3], and the tarsiers Tarsius lariang and T. dentatus [13]. Diagnosis Prior workers have recognized substantial species diversity in the genus Limnonectes on Sulawesi. However, diagnosing many of these lineages on the basis of morphology is challenging, and several authors have instead opted to apply names to Sulawesi specimens representing species from outside Sulawesi. Consequently, the following names have all been incorrectly applied to Sulawesi populations: the Lesser Sundas species (type locality: Flores Island) L. dammermani (Mertens, 1927), the Bornean species L. finchii (Inger, 1966), the Mollucan species (type locality: Ambon) L. grunniens (Daudin, 1801), and the Philippine taxa L. leytensis (Boettger, 1893), L. magnus (Stejneger, 1909), and L. palavanensis (Boulenger, 1894). These names should not be applied to Sulawesi populations, as was verified phylogenetically for several of these taxa [3]. Only four Sulawesi species have been described: L. arathooni (Smith, 1927), L. heinrichi (Ahl, 1933), L. modestus (Boulenger, 1882), and L. microtympanum (van Kampen, 1909). However, we will show elsewhere that L. heinrichi is a junior synonym of L. modestus and the species complex that we have referred to previously [2, 3] as L. modestus remains undescribed – thus, at present there are but three valid described species of Sulawesi Limnonectes. Limnonectes larvaepartus can be distinguished from all other described species of Limnonectes by its reproductive mode (Fig. 2). It can be further differentiated from all described Sulawesi species by its combination of body size (mean male SVL =37.4; female SVL = 40.2 mm), coloration, tympanum size, and degree of hind foot webbing, as well as on the basis of phylogenetic placement (Fig. 3). Limnonectes arathooni is endemic to Sulawesi’s Southwestern (SW) Peninsula south of the Tempe Depression, and thus does not occur within the range of L. larvaepartus. It is similar in size to the new species (male SVL = 36.6 mm, females= 39.6 mm), but differs in having substantially reduced webbing (extending to penultimate phalange of fourth toe vs. to toe disc), in lacking fine granular dorsal tubercles, and in having melanic spots above the forelimb insertion, a fine ridge extending posteriorly behind each eye, and an alternative derived reproductive mode in which males guard clutches of terrestrial eggs that hatch into tadpoles that then make their own way to an adjacent stream by sliding down steep stream-side embankments [14]. Limnonectes microtympanum, like L. arathooni, is restricted to the SW Peninsula south of the Tempe Depression, and thus does not overlap in geographical range with L. larvaepartus. Limnonectes microtympanum is moderately large (male SVL =78.4; female SVL = 72.4 mm), and thus much larger than the new species. Limnonectes microtympanum also differs from the new species in having proportionally smaller tympana (TY / SVL =0.05 + 0.01 in males, 0.06 + 0.01 in females versus 0.08 + 0.01 in both sexes in L. larvaepartus). The new species occurs in broad sympatry with L. modestus, which is a moderate sized (male SVL = 70.2 mm; female SVL =64.0 mm) inhabitant of fast-moving streams and substantially larger than L. larvaepartus. Like L. larvaepartus, L. modestus has nearly complete hindfoot webbing (slightly more extensive in L. modestus than in L. larvaepartus but reaching the toe disc in both species), a dusky throat with melanic pigments extending onto the pectoral region (in a clear wedge shape in L. modestus, more randomly distributed in L. larvaepartus), and skin with extensive fine granular tubercles. Limnonectes modestus also exhibits a derived reproductive mode involving production of a relatively small number of large (10 mm diameter) eggs that are deposited along the edge of fast moving streams. Description of the holotype An adult male (Fig. 1) 48 mm SVL, body moderately robust, head not broader than body, head about 65% longer than wide, length 45% of snout-vent length, snout 17% of snout-vent length, moderately pointed, projecting above lower jaw, nostril lateral, closer to tip of snout than to eye, lore essentially straight, canthus rostralis distinct, eye about equal to snout length, pupil diamond-shaped, upper eyelid with tubercles; interorbital region smooth, width 69% of internarial distance, tympanum moderate, slightly wider than interorbital distance, supratympanic fold distinct, extending from posterior corner of eye to supraaxillary region, in contact with tympanic annulus, temporal muscle slightly enlarged; odontoid process 2.1 mm. Dentigerous process of vomer distinct, angled anterolaterally, approximately at 45o angle, posterior ends separated by distance approximately equal to one-third diameter of choanae. Limbs relatively slender, tibia width at thickest part 7.5 mm; femur length 52% of snout-vent length, heels moderately overlapping when placed perpendicular to body axis; tibia length 64.9% of foot length, 48.9% of snout-vent length; foot length 71% of snout-vent length; tarsal fold indistinct, only evident as a ridge; toe discs moderately expanded, circum-marginal groove horseshoe-shaped, pointed anteriorly. Plantar surface of foot smooth, subarticular tubercle rounded, relative length of toes 4.3.5.2.1. Inner metatarsal tubercle prominent, elongate, ovoid with a sharp spade like ventral edge; outer metatarsal tubercle absent; hind foot webbing full, extending to toe discs, slightly emarginated. Manus length 51.3% of foot length, fingers slender, terminal discs slightly expanded, length formula 3.1$4$2, with slight differences in length, subarticular tubercle rounded, convex; supernumerary tubercles absent; inner and outer metacarpal tubercles enlarged, nuptial pads and webbing absent, forearm muscle not enlarged. See Table 1 for measurements and variation. Coloration The dorsal coloration is highly variable, typically brownish-grey, but can be darker brown on the dorsolateral region, and some individuals are reddish-brown or golden-tan (see Fig. 2). ~23% of specimens have a bold mid-dorsal stripe. The venter is either yellowish or cream colored, with the upper end of the tibia usually bearing a prominent dark spot. A light bar is often present in the interorbital region, and the coloration of the snout to interorbital region may be lighter than the remainder of the dorsum. The tympanum is often masked in black leaving only the lower rim sharing the predominant body coloration. The gular region is usually darker in males and may have a finely mottled wedge-shaped melanic patch. The dorsal half of the iris is golden-orange in coloration in at least some individuals (we do not know of exceptions, but have not documented iris coloration for most specimens). Limnonectes larvaepartus doi:10.1371/journal.pone.0115884.t001 Eggs and tadpoles Females produce ~100 non-pigmented eggs (see [15]), though the most we have observed is 55, which possibly represents the contents of just one oviduct. The eggs lack a jelly-coat and reach at least ~ 3 mm in diameter. These eggs develop within the oviducts into pigmented tadpoles that reach at least Gosner stage 35 prior to parturition (see [16] for staging). The gut is initially provisioned with substantial yolk (Fig. 2 b), and developing tadpoles prematurely removed from the oviducts at approximately stage 21 progressed to approximately stage 25 over the course of two weeks in a water bottle without supplemental food, suggesting plasticity in terms of the timing of parturition. A detailed description of the tadpole is provided in the accompanying paper by Kusrini et al. [15]. Natural history Limnonectes larvaepartus occurs in natural and disturbed forest habitats of Sulawesi, generally in sympatry with at least one, and sometimes as many as five other Limnonectes species. In the western Central Core of Sulawesi, we have always found L. larvaepartus living in sympatry with a much larger species that has been referred to in the literature as L. sp. D (2,3). Whereas L. sp. D is generally found on rocks in fast-moving streams or within a meter of water on the banks of fastflowing streams, L. larvaepartus is usually found further from the stream (2–10 meters from water) on rocky substrates, in leaf-litter, or secluded in grassy vegetation. Because we have observed that L. sp. D predates other frogs, including other Limonectes, it is possible that L. larvaepartus avoids large streams in response to predation pressure from larger Limnonectes species. Male L. larvaepartus typically call from the margins of seeps, puddles, or small pools away from the main stream. Notably, we have found many males calling from small pools that were already inhabited by L. larvaepartus tadpoles (Fig. 2 c), with as many as three size-classes of tadpoles represented. It is unclear whether some or all of the observed tadpoles were sired by the accompanying male. We have furthermore collected at least one pregnant female from a small stream-side puddle already inhabited by tadpoles of two size classes, again suggesting the possibility that individual pools may be visited repeatedly by the same adult males and females during the reproductive season.

Hemiphyllodactylus engganoensis sp. nov. Pulau Enggano Dwarf Gecko Cicak Kerdil Enggano Figs. 3, 4 Holotype. Adult female (MZB.Lace 4568) collected at 2220 hours on 11 May, 2013 by Jimmy A. McGuire, Djoko T. Iskandar, Awal Riyanto, and Mulyadi near the village of Malakoni on the island of Enggano (Kecamatan Enggano, Kabupaten Bengkulu, Propinsi Bengkulu, Indonesia; S 0 5.35290, E 102.27742, 1 m elevation). Paratype. Subadult female (MVZ 239345) and adult male (MVZ 239346) bear the same collection data as the holotype. Diagnosis. Hemiphyllodactylus engganoensis sp. nov. is differentiated from all other congeners by having the unique combination of a maximum SVL of 37.3 mm; six chin scales; no enlarged postmentals; five circumnasal scales; three or four scales between the supranasals; 12 supralabials; 24 or 25 dorsal scales; 14 ventral scales; a lamellar hand formula of 4554 or 4454; a lamellar foot formula of 4555; four or five subdigital lamellae on the first finger; four or five subdigital lamellae on the first toe; a contiguous femoroprecloacal pore series of 42; five cloacal spurs in males; no enlarged subcaudal scales; no dark postorbital stripes or striping on body; small dark blotches on dorsum; a yellowish postsacral mark bearing anteriorly projecting arms; and a pigmented caecum and gonads. (Table 1). Description of holotype. Adult female; head triangular in dorsal profile depressed, distinct from neck; lores and interorbital regions flat; rostrum relatively long (NarEye/ HeadL = 0.30); prefrontal region flat to weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate, rounded in dorsal profile; eye large; ear opening oval, small; eye to ear distance greater than diameter of eye; rostral wider than high, partially divided dorsally, bordered posteriorly by large supranasals; three internasals (=postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (collectively the circumnasals 5 R, L); 12 (R, L) square supralabials tapering to below posterior margin of orbit; 12 (R, L) square infralabials tapering to below posterior margin of orbit; scales of rostrum and lores, raised; scales on top of head and occiput small, granular; dorsal superciliaries raised, rectangular; mental triangular, bordered laterally by first infralabials and posteriorly by two non-enlarged postmentals; each postmental bordered laterally by a single sublabial; row of smaller scales extending transversely from juncture of second and third infralabials and contacting mental; gular scales triangular, small, granular, grading posteriorly into slightly larger, subimbricate throat and pectoral scales which grade into slightly larger, subimbricate ventrals. Body elongate, dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 25 scales contained within one eye diameter; ventral scales flat, subimbricate, much larger than dorsal scales, 14 scales contained within one eye diameter; no enlarged, precloacal scales; no pore-bearing scales femoral or precloacal scales; forelimbs short, robust in stature, covered with granular scales dorsally and slightly larger, flat, subimbricate scales ventrally; palmar scales flat, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal, subdigital lamellae of digits II���V divided, angular and U-shaped; lamellae proximal to these transversely expanded, undivided; lamellar formula of digits II���V 4 - 4 -5- 4 (R, L); four transversely expanded lamellae on digit I; claws on digits II���V well-developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; hind limbs short, more robust than forelimbs, covered with slightly pointed, juxtaposed scales dorsally and by larger, flat, subimbricate scales ventrally; plantar scales low, flat, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal, subdigital lamellae of digits II���V divided, angular and U-shaped; lamellae proximal to U-shaped lamellae transversely expanded, undivided; lamellar formula of digits II���V 4-5 - 5 - 5 (R, L); five transversely expanded lamellae on digit I; claws on digits II���V well-developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; all caudal scales flat, imbricate, not occurring in caudal segments, no enlarged subcaudals. Morphometric data are presented in Table 2. Coloration in alcohol. Top of head, body and limbs nearly unicolor tan; ground color of dorsal surface of tail tan bearing nine, dark-colored, diffuse bands; lores slightly darker; no postorbital striping; faint, diffuse lightcolored spots on dorsum barely discernable; light-colored, postsacral marking bearing faint, anteriorly projecting arms; ground color of gular region beige with small, dark-brown spots; ground color of ventral surfaces of body and limbs beige, immaculate; dorsal coloration invades lateral sections of abdomen. Variation. The paratype MVZ 239345 approaches the holotype in general dorsal coloration and pattern (Fig. 3, 4). In life, the dorsal ground color of the head is dull-yellow and that of the body, limbs, and tail is tan with faint, darker bands. All dorsal surfaces are overlain with a reticulum of darker markings that tend to form thin, zig-zag lines across the body and wider bands on the tail. The light-colored, postsacral marking is more vivid. The iris is silver. The tail is complete, original, and round in cross-section (Fig. 4). The male paratype MVZ 239346 is missing a tail and has a uniform tan dorsal ground color (Fig. 4). It also has a continuous series of 42 femoroprecloacal pores and five cloacal spurs, both of which are lacking in the two female specimens. Differences in meristics and morphometrics are listed in Table 2. MZB.Lace MVZ MVZ Distribution. Hemiphyllodactylus engganoensis sp. nov. is known only from the type locality near the village of Malakoni on the island of Enggano. We presume it to be endemic to the entire island, though it may be restricted to Enggano���s coastal perimeter. All three specimens in the original series were collected within a few hundred meters of one another in beachside vegetation. Natural history. The three specimens of Hemiphyllodactylus engganoeneis sp. nov. were collected where lowland forest habitat interfaced with a sandy beach near the mouth of a small river along the Enggano coastline. The holotype was collected approximately 1 m above the ground on the leaf of a sapling, and the two paratypes were collected about 1 m above the ground on and between Pandanus leaves. The specimens were all collected between 2200 and 2225 hrs. Pandanus was abundant along the forest edge, and the crevices between its serrated, strap-shaped leaves were also inhabited by two other gecko species, Lepidodactylus lugubris (Dum��ril & Bibron) and Hemidactylus frenatus Schlegel. Etymology. The specific epithet engganoensis is an adjective in reference to the type locality Pulau Enggano, Bengkulu Province, Indonesia. Comparisons. The taxonomy of Grismer et al. (2013, 2014), Ngo et al. (2014); Nguyen et al. (2013), and Zug (2010) is used in the comparisons below for H. titiwangsaensis Zug, H. typus Bleeker and H. yunannensis (Boulenger). Hemiphyllodactylus engganoensis sp. nov. is one of the smallest species of the genus and differs from H. banaensis Ngo, Grismer, Thai, & Wood; H. chiangmaiensis Grismer, Wood, & Cota; H. larutensis (Boulenger); H. margarethae Brongersma; H. titiwangsaensis Zug; H. typus Bleeker; H. yunnanensis; and H. zugi Nguyen, Lehmann, Le Duc, Bonkowski, & Ziegler by its maximum SVL of 37.3 mm vs a SVL>41.0 mm. Its postmentals are not distinctly enlarged which separates it from H. banaensis, H. chiangmaiensis; H. harterti (Werner); H. larutensis; H. margarethae; H. titwangsaensis; H. typus; H. yunnanensis; H. tehtarik Grismer, Wood, Anuar, Muin, Quah, McGuire, Brown, Ngo, & Thai; and H. zugi. Hemiphyllodactylus engganoensis sp. nov. can be differentiated from H. banaensis, H. larutensis, H. titwangsaensis, and H. zugi by having five vs. two or three circumnasal scales. It differs from all other species except H. zugi in having 24 or 25 dorsal scales (as opposed to having fewer than 22) and having 14 ventral scales separates it from H. banaensis, H. aurantiacus (Beddome), H. chiangmaiensis, H. ganoklonis Zug, H. margarethae, H. titwangsaensis, H. yunnanensis, and H. tehtarik which have 12 or less. Hemiphyllodactylus engganoensis sp. nov. has a unique lamellar formula on the hand (4-5 - 5 - 4 or 4 - 4-5 - 4) that separate if from all other species except H. banaensis that have various other combinations (see Table 1). Having a series of femoral and precloacal pores that are not contiguous separates H. engganoensis sp. nov. from H. aurantiacus, H. ganoklonis, H. insularis Taylor, H. margarethae, some H. typus, and H. yunnanensis which all have a contiguous pore series. Additionally, having a total pore count of 42 further differentiates H. engganoensis sp. nov. from all other species (except H. harterti) who have less than 40 (0���39 collectively). Hemiphyllodactylus engganoensis sp. nov. has a high number (5) of cloacal spurs which at least distinguishes it from H. auratiacus, H. banaensis, H. harterti, H. insularis, H. larutensis, H. margarethae, H. yunnanaensis, and H. zugi which collectively have 0���3 cloacal spurs. There are a number of morphometric ratios concerning various structures that are potentially diagnostic (Table 1). However, due to the incomplete sample sizes across all age classes for all species, we consider their diagnostic utility as tentative at this point. Additionally, H. engganoensis sp. nov. is possibly separated from various other species on the basis of a number color pattern characters (Table 1). However, because the color variation in the type series (three specimens) is extensive (Fig. 3), we defer judgment as to the utility of these characters as well until the acquisition of additional material. Based on the mitochondrial ND 2 gene, Grismer et al. (2013) noted that H. engganoensis sp. nov. has an uncorrected sequence divergence of 17.5 % from its closest relative from Pulau Sibu, Malaysia., Published as part of Grismer, L. Lee, Riyanto, Awal, Iskandar, Djoko T. & Mcguire, Jimmy A., 2014, A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Pulau Enggano, southwestern Sumatra, Indonesia, pp. 485-495 in Zootaxa 3821 (4) on pages 487-494, DOI: 10.11646/zootaxa.3821.4.7, http://zenodo.org/record/228521, {"references":["Ngo, V. T., Grismer, L. L., Thai, P. H. & Wood, P. L. (2014) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Ba Na-Nui Nature Reserve, central Vietnam. Zootaxa, 3760 (4), 539 - 552. http: // dx. doi. org / 10.11646 / zootaxa. 3760.4.3","Nguyen T., Lehmann T., Le M. D., Duong H. T., Bonkowski, M. & Ziegler, T. (2013) A new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Vietnam. Zootaxa, 3736 (1), 89 - 98. http: // dx. doi. org / 10.11646 / zootaxa. 3736.1.5","Zug, G. R. (2010) Speciation and dispersal in a low diversity taxon: the Slender geckos Hemiphyllodactylus (Reptilia, Gekkonidae). Smithsonian Contributions to Zoology, 631, 1 - 70."]}

Grismer, L. Lee, Riyanto, Awal, Iskandar, Djoko T., Mcguire, Jimmy A. (2014): A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Pulau Enggano, southwestern Sumatra, Indonesia. Zootaxa 3821 (4): 485-495, DOI: http://dx.doi.org/10.11646/zootaxa.3821.4.7

Penelitian ini bertujuan untuk mengetahui ada tidaknya polimorfisme gen pengkode membran peritrofik PM-48 asal Bandung dan Makassar. Membran peritrofik dihasilkan oleh kardia larva Crysomya bezziana dan merupakan suatu kandidat vaksin yang ideal untuk mencegah penyakit miasis pada sapi. Telah dilakukan penelitian tentang polimorfisme gen pengkode protein membran peritrofik PM-48 Chrysomya bezziana yang berasal dari Bandung dan Makassar. Tujuan penelitian ini adalah untuk membandingkan gen pengkode membran peritrofik PM-48 asal Bandung dan Makassar. Membran peritrofik dihasilkan oleh kardia larva Crysomya bezziana dan merupakan suatu kandidat vaksin yang ideal untuk mencegah penyakit miasis pada sapi. Salah satu dari protein membran peritrofik tersebut adalah PM-48, yang telah berhasil diklon dan diurutkan pasangan basanya. Metode yang digunakan adalah isolasi DNA genom, mengamplifikasi dengan PCR menggunakan primer spesifik SW PM-48. Selanjutnya produk PCR dipotong dengan enzim restriksi Hinf I, Kpn I dan EcoR I. Hasil ampifikasi dengan PCR pada 60 sampel diperoleh fragmen berukuran 1,2 kb. Setelah dilakukan pemotongan dengan Hinf I diperoleh tiga larik ukuran 0,1 kb, 0,3 kb, dan 0,8 kb sedangkan dengan Kpn I dihasilkan dua larik dengan ukuran 0,3 kb dan 0,9 kb. Pemotongan dengan EcoR I menghasilkan dua larik dengan ukuran 0,5 kb dan 0,7 kb. Dari pola restriksi yang dihasilkan setiap individu lalat, diketahui bahwa tidak terdapat polimorfisme gen pengkode protein membran peritrofik Chrysomya bezziana yang berasal dari Bandung dan Makassar

Iskandar, Djoko T., Rachmansah, Angga (2011): A new bent-toed gecko of the genus Cyrtodactylus Gray, 1827 (Reptilia, Gekkonidae) from Mount Tompotika, eastern peninsula of Sulawesi, Indonesia. Zootaxa 2838: 65-78, DOI: 10.5281/zenodo.206737

Cyrtodactylus batik sp. nov. (cicak batik; batik bent-toed gecko) (Figs. 1 A, B; 2; 3 A; 4 A) Holotype. ITB. DTI 2805, an adult female with original tail from Longkoga Stream, Bualemo, Mount Tompotika, Balantak Mountains (between 00�� 40 '05.1"��� 00�� 40 ' 12.7 "S; 123 ��06' 41.7 "��� 123 ��06' 39.2 "E; alt: 951���1002 m asl), Desa Trans Tanah Merah, Kecamatan Bualemo, Kabupaten Banggai, Propinsi Sulawesi Tengah, Sulawesi Island, Indonesia collected by A. Rachmansah and Umilaela on 20 May 2009. Paratypes. DTI 2784, DTI 2801, DTI 2804, DTI 2785, DTI 2803, DTI 2802, same data as for the holotype, collected 19���20 May 2009. Diagnosis. A large form of Cyrtodactylus with SVL reaching 115 mm in adult females, males slightly smaller, up to 110 mm, tail 108���120 % of SVL; body robust, limbs medium length; digits long; single pair of postmentals in contact posteriorly, isolating triangular mental from chin shields; dorsum with 23���26 transverse rows of slightly keeled trihedral tubercles, slightly larger than adjacent dorsal scales giving a generally smooth appearance; 48���57 smooth, round, juxtaposed ventral scales between distinct ventrolateral folds; no precloacal groove, no precloacal or femoral pores, no enlarged femoral scales; distinctly enlarged precloacal scale patch; 7���10 transversely expanded lamellae proximal to basal inflection of 4 th toe, 10���16 narrow lamellae distal to inflection. Underside of the hemipenal bulge of tail base bearing approximately 30 rows of small postcloacal scales, followed by approximately five rows of slightly enlarged, rectangular subcaudals followed by transversely expanded subcaudals. Etymology. The specific epithet is used as a noun in apposition, originating from the specific Indonesian pattern of traditional ��� batik ��� cloth that is especially well known on Java. The dorsal pattern of the new species is similar to that of traditional batik cloth. Holotype description. An adult female, SVL 103.2 mm (104.8 mm measured prior to fixation after being euthanized), TL 115.1 mm (fresh 118.2 mm, TL/SVL ratio 1.08���1.20). Head moderately long (HL/SVL ratio 0.29), wide (HW/HL ratio 0.60), moderately depressed (HH/HL ratio 0.34), distinct from neck. A raised, rounded supraorbital ridge continuous with canthus rostralis. Distinct frontoparietal depressions posterior to each supraorbital prominence. Lores weakly convex anteriorly, mildly depressed posteriorly; separated from anterior palpebrals and orbits by deep lacrimal grooves. Dorsal surface of snout anteriorly swollen above nostrils. Lacrimal groove met orthogonally by midpalpebral depression continuing around orbit. Small, raised extension of skin comprised of superciliaries and distalmost rows of palpebrals extending to the circumference of the orbit. Supraorbital scales uniform, lacking small tubercles. Superciliaries large, composed of two rows of overlapping scales. These scales are relative long, uniform, height ranged from 0.9 mm at the sides to 1.2 mm above the center of eye; forming a crenulated, erect rim around eye, rounded and smooth, without distinct keel, 16���21 rows of interorbital tubercles across narrowest point of frontal bone. Snout relatively short (SL/HL ratio 0.40); longer than eye diameter (OD/SL ratio 0.59). Scales of snout round, granular, uniform in size, rostrum scales smooth, without tubercles; tiny tubercles present on medial palpebral and interorbital regions. Head tubercles pointed, symmetrical, larger posteriorly, attaining maximum size at occiput. Scales of rostrum granular, regular in size (as in snout region). Eyes moderate (OD/HL ratio 0.23). Auricular openings erect, egg shaped, narrower dorsally and rounded ventrally; openings large (EaL/HL ratio 0.14); eye to ear distance slightly greater than diameter of eye (EaEy/OD ratio 1.1). Rostral 50 % deeper (2.1 mm) than wide (4.2 mm) at narrowest point, 69 % as deep (2.9 mm) as wide (4.6 mm) at longest point; incompletely divided by dorsal Y��� shaped rostral groove; two supranasals, anteriormost pair separated by pair of median postinternasals and two internasal scales; rostral in contact with first supralabial, anteriormost supranasal, median postinternasal and internasal. Nares oval, oriented laterally, in contact with two supranasals, first supralabial; 3 / 3 postnasals, about the same size of head or body granular scales; Mental triangular in shape, nearly twice as wide as deep; single pair of postmentals contacting posteriorly about half their length. Postmentals bordered laterally by first infralabials, the anterior tips extend to the suture between mental and first sublabials; posteriorly by two slightly enlarged chin shields (adjacent to second infralabial), 2 intermediate sized and seven small irregularly shaped, granular gular scales. Throat covered with uniform granular scales. Nine supralabials to midpoint of orbit (13 to angle of jaw). Eight infralabials to midpoint of orbit (12 on the left and 11 on the right to angle of jaw). Body slender (TrL/SVL ratio 0.49); indistinct ventrolateral folds with about a dozen rounded, white tubercles. Dorsum between forelimb insertion and caudal region characterized by small granular scales interspersed with irregularly spaced, keeled unicarinate tubercles. Anterior side of tubercles rounded, rising gradually, defined by a single pronounced median, rounded keel; posterior sides convex, steeply sloping. From forelimb insertion to frontal region, tubercles progressively decrease in size, becoming round, non-keeled on head. A total of 36 paravertebral tubercles between forelimb and hind limb insertions are present. Ventral scales smooth, juxtaposed, regular in size; ventrals larger in diameter than both dorsal granular scales and some dorsal tubercles; gulars small, granular, uniform; 49 scales between dorsolateral folds across midsection of body. No precloacal pores or precloacal groove. No femoral pores or enlarged femoral scales, but enlarged precloacal scales present. Scales adjacent the precloacal area and on the lower parts of the femur become abruptly smaller compared to those at the precloacal region. Limbs medium in length, moderately robust; forearms shorter than hind limbs; forearm short (FaL/SVL ratio 0.17); tibia short (CrL/ SVL ratio 0.18); suprabrachials and prebrachials larger than scales of adjacent dorsum, granular, tuberculate, much smaller in postbrachial region proximal to elbow; infrabrachials small, granular, about equal to gulars; postantebrachials granular at elbow; numerous tuberculation present on tibial surfaces, except infratibial surfaces; supraantetibials regular, granular proximally, becoming large, continuous, unchanged in size with supracarpals and supradigital lamellae; infraantebrachial squamation similar to supraantebrachials, though tubercles not present; suprafemorals and prefemorals similar to supratibials, scales regular, smooth, tubercles numerous, well differentiated compared to neighboring granules; infrafemorals small, similar to granules on adjacent body; no enlarged infrafemoral or postfemoral scale series, squamation similar to adjacent interfemorals; ventrals, supratibials and pretibials small, granular, as in ventral femoral regions; unlike supracarpal squamation, supratarsals consistently small, granular, met abruptly by enlarged supradigital lamellae; infratarsals similar in size proximally, but becoming enlarged, subimbricate distally. Digits long, strongly inflected at basal interphalangeal joints. Claws large (maximum length of 1.9 mm), surrounded by elongate, deeply notched distal-most subdigital lamella, one enlarged supradigital scale, and a smaller (about half size) lateral scale. Subdigital lamellae elongate, narrow distal to first interphalangeal joint. Subdigital lamellae proximal to first interphalangeal joint wider than long, swollen, padlike, especially at interphalangeal joint. Counts of subdigital lamellae on manus I: 15 / 15; II: 18 / 19; III: 20 / 20; IV: 21 / 22; V: 19 / 19, on pes I: 15 / 14; II: 19 / 19; III: 23 / 23; IV: 24 / 26; V: 23 / 24; webbing absent. Relative lengths of digits on manus: IV> III> II> I> V, on pes: IV> V> III> II> I. Tail relatively long, (TL/SVL ratio 1.11); portion of tail subrectangular at base with regularly���spaced strongly trihedral, keeled tubercles more sparsely distributed than tubercles of dorsum; three prominent, enlarged post���cloacal spurs on each side of vent; subcaudals arranged in several rows of small, narrow rectangular scales followed by enlarged median subcaudal plates variable in size, largest 3.9 mm wide, 1.2 mm long; most approximately half this size. Holotype coloration. Overall dorsal appearance uniform velvety black, tubercles have the same color as background hence not visible on photographs except on the flanks. Four pairs of overlapping ��� > shaped irregular yellow transverse bands between nape and base of tail and 10 similar marks on the tail, the first three ��� >< ��� shaped marks on the tail are more or less similar to the dorsal pattern, the remainder less distinct in form with yellow spots and crosses. The areas within the overlapping ��� > shaped marks are lighter compared to the dorsum. A yellow line borders the posterior margin of the head. Limbs with irregular yellow bands or spots at various angles; distinct yellow bars at the finger-carpal joint; head coloration slightly lighter than dorsum, faintly marbled with yellow spots which are variable in size, a yellow line running along superciliaries to occiput enclosing parietal region of head and posterior part of canthus rostralis; upper labials scales generally lighter with some yellow spots along the upper border; rostral as dark as body coloration; nape dark; eyes essentially black, iris greenish metallic during daylight (see Fig 1 B); lateral surfaces similar to dorsum but with yellow tubercles, sparsely arranged on the flanks, strongly contrasted with velvety black base color; venter and undersides of limbs uniformly blackish, ventral scales with numerous fine purple flecks covering otherwise pale scales. Variation in the paratypes. The head varies slightly in length (HL/SVL ratio 0.29���0.32), width (HW/HL ratio 0.59���0.65), and moderately depressed (HH/HL ratio 0.27���0.34); interorbital tubercles across narrowest point of frontal bone varies between 16���21 rows. Snout relatively short (SL/HL ratio 0.39���0.43); longer than eye diameter (OD/SL ratio 0.56���0.58). Eyes moderate (OD/HL ratio 0.22���0.25). Auricular openings is large (EaL/HL ratio 0.12���0.14); eye to ear distance slightly greater than diameter of eye (EaEy /OD ratio 1.1���1.3). Rostral varies from 43���53 % deeper (1.8���2.1 mm) than wide (4.2���4.6 mm) at narrowest point, 67���88 % as deep (2.9���3.7 mm) as wide (4.6���5.6 mm) at longest point. Anteriormost pair of SuN separated by a pair of median post InNs and two to three InNs. PM bordered laterally by first and/or second InLs, the anterior tips extend to the suture between mental and first InLs; posteriorly by two slightly enlarged chin shields (adjacent to second InL) and two to eight intermediate sized and up to seven small irregularly shaped, granular gular scales. Supralabials varies from nine or ten to midpoint of orbit (13���15 to angle of jaw). Body slender (TrL/SVL ratio 0.39���0.49). A total of 33���40 PVTs between forelimb and hind limb insertions are present. Ventrals ranges from 48���57 scales between dorsolateral folds across midsection of body. The limbs are medium in length, forearm short (FaL/SVL ratio 0.15���0.17); tibia short (CrL/ SVL ratio 0.17���0.19); Variations of subdigital lamellae on manus I: 13���16; II: 18���20; III: 20���23; IV: 21���23; V: 18- 22, on pes I: 14���16; II: 19���21; III: 23���25; IV: 24��� 27; V: 20���24; webbing absent. Tail relatively long, (tail length/ SVL ratio 1.08���1.20). For other detailed measurements and detailed count of the whole type series see Table 1. Regenerated tail completely round; caudals of re���grown tail extremely reduced, tubercles absent; subcaudals of regenerated part of tail in proportion to original subcaudals, but reduced in size and shorter. Scales of regrown tail similar in shape, lightly colored, mottled, lacking tubercles and the ��� >< ��� shaped marks (see Fig 1 B). Secondary sex characteristics. From specimens on hand, the male attains a size nearly as large as females, and can be distinguished by the presence of bulging hemipenes at the base of the tail, whereas females have a slen- der tail base. Both sexes lack precloacal and femoral pores, and hence these are not useful for determining sex of this species. Ecological notes. All specimens were found on vegetation in undisturbed primary forest, from 1.5 to 3 meters above ground and more than 50 meters from the closest stream (Longkoga) in primary forest. The trees format the collecting sites were lianas and small trees with trunks less than 40 cm in diameter. The smaller species, C. jellesmae was not found in the same site, but at other camp sites it occurred approximately 1���2 meters above the ground on tree trunks and smaller vegetation. Comparison between species. Cyrtodactylus batik and C. wallacei have a similar coloration pattern (Fig 2 A, B), but the basic coloration of C. batik is black with yellow (cream) transverse stripes, sharply demarcated with tubercles having the same coloration as the background except on the flanks. In C. wallacei, the dorsal ground coloration is reddish brown with weakly defined lighter stripes made up by light colored tubercles on the dark background. The scales and tubercles of C. batik are smaller compared to those of C. wallacei. Consequently, every detail in number of scales, head scales, tubercles as well as number of lamellae under finger and toes are higher in C. batik, except for the number of first and second toe lamellae, which are higher in C. wallacei (see Table 2). Despite similarity in size, the banding pattern, velvety black dorsum, differences in number of tubercles, and number of ventral scales distinguish it from the very similar and geographically close C. wallacei. The smaller size and light brown dorsal coloration with sharply defined blotching pattern distinguish C. jellesmae from C. batik. These three Sulawesi species are similar in lacking precloacal and femoral pores. Based on examination of gravid females, C. jellesmae is a species complex with at least two different body size groups (Table 2). The smaller form is widely distributed, has fragmented subcaudals and shares the presence of tubercles along the ventrolateral fold with C. spinosus and C. batik. The larger form is restricted to southeast Sulawesi and is characterized by the absence of enlarged tubercles along the ventrolateral fold, but has broad transverse subcaudals as in C. batik. Otherwise both forms currently identified as C. jellesmae have similar dorsal coloration (aside for other small forms with different dorsal blotching pattern and not included in the analysis). Cyrtodactylus batik is distinguishable from all but eight congeneric species by the absence of a precloacal groove, and precloacal and femoral pores. The first two (three if the large form identified as C. jellesmae is counted as a separate species) occur in Sulawesi and have been discussed previously. The six remaining congeners may be distinguished from C. batik by the following characteristics: tiny dorsal conical tubercles, a black dorsal coloration and large size (maximum SVL 113 mm) distinguish C. batik from C. laevigatus (Darevsky, 1964; presence of an enlarged femoral scale row forming a distinct boundary and separating smaller posterior femorals distinguish C. paradoxus (Darevsky & Szczerbak, 1997); presence of enlarged femoral scales, smaller SVL and dark bands or blotches contrasting with a light gray background distinguish C. semenanjungensis (Grismer & Leong, 2005); presence of a quadrangular rostral bordered by fewer scales and with a single median cleft, smaller size, and heavy yellow spotting on arms, dorsum and, most distinctively, along ventrolateral folds and labial regions distinguish C. sermowaiensis (de Rooij, 1915); presence of an extremely enlarged femoral and precloacal scale series, slightly smaller size and distinct reddish-orange and black bandings distinguish C. thirakhupti (Pauwels et al.). Cyrtodactylus malayanus (de Rooij, 1915) and C. consobrinus (Peters, 1871) are two species from Borneo, about equal in size, that differ dramatically from the Sulawesi forms by their banded color pattern and in having enlarged femoral scales. Precloacal and femoral pores may be present in some individuals, but these, in combination with different dorsal coloration, set them apart from all Sulawesi forms. Cyrtodactylus spinosus and Cyrtodactylus sp. from Sulawesi Barat (see Appendix 1) both have precloacal pores, while C. fumosus has both precloacal and femoral pores in a continuous series. The lack of spines along the ventrolateral flanks and head, as well as size and blotching pattern distinguish C. batik from C. spinosus. Sulawesi. *) Data on C. wallacei and C. spinosus were extracted from the original description. n.a = data not available. In the Lesser Sunda Islands three other species have been described: C. darmandvillei (Weber, 1890), C. gordongekkoi (Das, 1993) and C. wetariensis (Dunn, 1927). Each has precloacal and femoral pores, hence they are easily distinguished from C. batik. From Maluku, three species have been recognized, C. halmahericus (Mertens, 1929) from Halmahera and Seram; C. deveti (Brongersma, 1948) from Morotai Island and C. nuaulu (Oliver et al., 2009) from Seram Island. As all three species have precloacal and femoral pores, they are easily distinguished from C. batik. Regarding coloration differences, C. deveti has bold bars on the dorsum and C. nuaulu is ornamented with elongated bands along paravertebral area, hence these are easily distinguishable from C. batik. Cyrtodactylus halmahericus has a number of poorly defined bands and hence is also easily distinguishable from C. batik. It is interesting that among the eight poreless species mentioned above, four occur in the Wallacean region, three are present in Southeast Asia and one in New Guinea. The absence of precloacal pores, femoral pores and an enlarged femoral scale series in C. batik, C. wallacei, C. jellesmae complex, C. laevigatus and C. sermowaiensis suggests these species may be closely related. Three species are Sulawesi endemics. In contrast, C. laevigatus is endemic to Lesser Sunda Islands while C. sermowaiensis is restricted to the north coast of New Guinea. Nothing is known about the evolutionary process towards pore loss or other frequently-used diagnostic morphological characters in Cyrtodactylus. Change in body size and the wide transverse subcaudal scales are plausible paths of evolutionary change. Several other species without precloacal and femoral pores have been found in Sumatra, but these forms, Published as part of Iskandar, Djoko T. & Rachmansah, Angga, 2011, A new bent-toed gecko of the genus Cyrtodactylus Gray, 1827 (Reptilia, Gekkonidae) from Mount Tompotika, eastern peninsula of Sulawesi, Indonesia, pp. 65-78 in Zootaxa 2838 on pages 66-74, DOI: 10.5281/zenodo.206737, {"references":["Darevsky, I. S. (1964) Two new species of gekkonid lizards from the Komodo Island in Lesser Sundas Archipelago. Zoologischer Anzeiger, 173, 169 - 174.","Darevsky, I. S. & Szczerbak, N. N. (1997) A new gecko of the genus Gonydactylus (Sauria: Gekkonidae) with a key to the species from Vietnam. Asiatic Herpetological Research, 7, 19 - 22.","Grismer, L. L. & Leong, T. M. (2005) New species of Cyrtodactylus (Squamata: Gekkonidae) from Southern Peninsular Malaysia. Journal of Herpetology, 39, 584 - 591.","de Rooij, N. (1915) The Reptiles of the Indo-Australian Archipelago. I. Lacertilia, Chelonia, Emydosauria. E. J. Brill Ltd., Leiden.","Peters, W. C. H. (1871) Uber neue Reptilien aus Ostafrica und Sarawak (Borneo), vorzuglich aus der Sammlung des Hrn. Marquis J. Doria zu Genoa. Monatsbericht Koniglich Preussischen Akademie der Wissenschaften zu Berlin, 1871, 566 - 581.","Weber, M. C. W. (1890) Reptilia from the Malay Archipelago. 1. Sauria, Crocodylidae, Chelonia, pp 158 - 177. In M. C. W. Weber (Ed.) Zoologische Ergebnisse einer Reise in Niederlandisch ost - Indien, I. E. J. Brill, Leiden.","Das, I. (1993) Cnemaspis gordongekkoi, a new gecko from Lombok, Indonesia, and the biogeography of oriental species of Cnemaspis (Squamata: Sauria: Gekkonidae). Hamadryad, 18, 1 - 9.","Dunn, E. R. (1927) Results of the Douglas Burden expedition to the island of Komodo III. Lizards from the East Indies. American Museum Novitates, 288, 1 - 13.","Mertens, R. (1929) Zwei neue Haftzeher aus dem Indo-Australischen Archipel (Rept.). Senckenbergiana Biologica, 11, 237 - 241.","Brongersma, L. D. (1948) Lizards from the island of Morotai (Moluccas). Proceedings of the Koninklijke Nederlandsche Akademie van Wetenschappen, Amsterdam, 51, 486 - 495.","Oliver, P., Edgar, P., Mumpuni, Iskandar, D. T. & Lilley, R. (2009) A new species of bent-toed gecko (Cyrtodactylus: Gekkonidae) from Seram Island, Indonesia. Zootaxa, 2115, 47 - 55.","Hall, R. (1996) Reconstructing Cenozoic SE Asia. In R. Hall & D. Blundell (Eds.), Tectonic Evolution of Southeast Asia. Vol. No. 106. Geological Society of London Special Publications, London, U. K. Pp. 153 - 184.","Hall, R. (1998) The plate tectonics of Cenozoic SE Asia and the distribution of land and sea. In R. Hall & J. D. Holloway (Eds.), Biogeography and Geological Evolution of SE Asia. Backhuys Publishers, Leiden, The Netherlands. Pp. 99 - 131.","Hall, R. (2001) Cenozoic reconstructions of SE Asia and the SW Pacific: Changing patterns of land and sea. In I. Metcalfe, et al. (Eds.), Faunal and Floral Migrations and Evolution in SE Asia-Australasia. A. A. Balkema (Swets & Zeitlinger Publishers), Lisse, The Netherlands. Pp. 33 - 56.","Moss, S. J. & Wilson, E. J .. (1998) Biogeographic implications of the Tertiary palaeogeographic evolution of Sulawesi and Borneo. In Hall, R. & Holloway, J. D. (Eds.), Biogeography and Geological Evolution of SE Asia. Backhuys Publishers, Leiden, The Netherlands. Pp. 133 - 163.","Whitten, A. & Whitten, J. (1992) Wild Indonesia. World Wildlife Fund for Nature, New Holland, London, U. K.","Whitten, A. J., Damanik, S. J., Anwar, J. & Hisyam, N. (2000) The Ecology of Sumatra. Periplus Editions, Singapore.","How, R. A. & Kitchener, D. J. (1997) Biogeography of Indonesian Snakes. Journal of Biogeography, 24. 725 - 735","Iskandar, D. T., & Tjan, K. N. (1996) The amphibians and reptiles of Sulawesi, with notes on the distribution and chromosomal number of frogs. In Kitchener, D. J. & Suyanto. A. (Eds.), Proceedings of the First International Conference on Eastern Indonesian- Australian Vertebrate Fauna. Manado, Indonesia. Pp. 39 - 46.","Iskandar, D. T. & Colijn, E. (2000) Preliminary checklist of Southeast Asian and New Guinean Herpet (o) fauna I. Amphibians. Treubia, 31, 1 - 134.","Iskandar, D. T. & Colijn, E. (2001) A Checklist of Southeast Asian and New Guinean Reptiles I. Serpentes. Biodiversity Conservation Project, (Indonesian Institute of Sciences), Japan International Cooperation Agencies, Ministry of Forestry, Institut Teknologi Bandung and The Gibbon Foundation. Jakarta, Binamitra. 195 pp.","Inger, R. F. & Voris, H. K. (2001) The biogeographical relations of the frogs & snakes of Sundaland. Journal of Biogeography. 28, 863 - 891.","McGuire, J. A., Brown, R. M., Mumpuni, Riyanto, A. & Andayani, N. (2007) The flying lizards of the Draco lineatus group (Squamata: Iguania: Agamidae): A taxonomic revision with descriptions of two new species. Herpetological Monographs, 21, 180 - 213.","Shekelle, M., Groves, C., Merker, S. & Supriatna, J. (2008) Tarsius tumpara: A new tarsier species from Siau Island, North Sulawesi. Primate Conservation, 23, 55 - 64.","Gressit, J. L. (1961) Problems in zoogeography of Pacific and Antarctic insects. Pacific Insects Monographs, 2, 1 - 94.","Whitten, A. J., Mustafa, M. & Henderson, G. S. (1987) The Ecology of Sulawesi. Gadjah Mada University Press, Yogyakarta, Indonesia.","Evans, B. J., Brown, R. M., McGuire, J. A., Supriatna, J., Andayani, N., Diesmos, A., Iskandar, D. T., Melnick, D. V. & Cannatella, D. (2003 a) Phylogenetics of fanged frogs: testing biogeographical hypotheses at the interface of the Asian and Australian faunal zones. Systematic Biology, 52, 794 - 819.","Evans, B. J., Supriatna, J., Andayani, N., Setiadi, M. I., Cannatella, D. C. & Melnick, D. J. (2003 b) Monkeys and toads define areas of endemism on Sulawesi. Evolution, 57, 1436 - 1443.","Howard, S. D., Gillespie, G. R., Riyanto, A. & Iskandar, D. T. (2007) A new species of large Eutropis (Scincidae) from Sulawesi, Indonesia. Journal of Herpetology, 41, 604 - 610.","Brown, R. M., Supriatna, J. & Ota, H. (2000) Discovery of a new species of Luperosaurus (Squamata; Gekkonidae) from Sulawesi, with a phylogenetic analysis of the genus and comments on the status of L. serraticaudus. Copeia, 2000, 191 - 209.","Inger, R. F. & Marx, H. (1965) The systematics and evolution of the Oriental colubrid snakes of the genus Calamaria. Fieldiana Zoology, 49, 1 - 304.","Koch, A., Arida, E., McGuire, J. A., Iskandar, D. T. & Bohme, W. (2009) A new species of Calamaria (Squamata: Colubridae) similar to C. ceramensis de Rooij, 1913, from the Banggai Islands, east of Sulawesi, Indonesia. Zootaxa, 2196, 19 - 30."]}

Key to species of Cyrtodactylus of Sulawesi This key is mainly based on morphological characteristics of adult males. 1 a. Precloacal groove and pores present in males................................................................ 2 1 b. Precloacal groove absent in males and females............................................................. 4 2 a. A medium sized species, 42–56 precloacal-femoral pores, sometimes a number of infrascales present in males (North Sulawesi)................................................................................... C. fumosus 2 b. Medium to large species, femoral pores lacking, males with precloacal pores...................................... 3 3 a. Precloacal pores 12–14 in males, spines along ventrolateral body fold and ventrolateral margin of tail (Lore Lindu, Central Sulawesi)................................................................................... C. spinosus 3 b. Precloacal pores 9–12 in males, no spines along ventrolateral body fold and ventrolateral margin of tail (West Sulawesi).......................................................................................... Cyrtodactylus sp. 4 a. No precloacal or femoral pores; SVL less than 93 mm........................................................ 5 4 b. No precloacal or femoral pores; SVL greater than 92 mm...................................................... 6 5 a. SVL 73–93 mm, subcaudals composed of transverse widened scales (Southeast Sulawesi)............. C. jellesmae large 5 b. SVL 58–70 mm, subcaudals scales fragmented (widespread allover Sulawesi)....................... C. jellesmae small 6 a. Dorsum brownish purple with distinct light colored tubercles and four overlapping “ >< ” shaped marks composed of light col- ored tubercles (West Sulawesi and Kabaena)........................................................ C. wallacei 6 b. Dorsum velvety black with four overlapping “ >< ” shaped marks of yellow stripes (Eastern Peninsula of Sulawesi)... C. batik

While land-use change is rapid throughout Southeast Asia, the island of Sulawesi (Indonesia) is of pressing conservation concern because of its exceptional number of endemic species. However, a lack of good identification literature for certain taxa such as amphibians and reptiles (apart from snakes) substantially delays ecological research in this region. Here, we compile an illustrated species list based on three years of research in and around the Lore Lindu National Park (LLNP) area and supplement it with data from the literature. In total, our survey and the literature review revealed 25 amphibian and 54 reptile species in five and 13 families, respectively. Our results highlight the LLNP area as an important herpetological endemism hotspot in the region. Appropriate utilization of species lists like this may facilitate capacity-building of local scientists and knowledgable local guides working in ecotourism. While land-use change is rapid throughout Southeast Asia, the island of Sulawesi (Indonesia) is of pressing conservation concern because of its exceptional number of endemic species. However, a lack of good identification literature for certain taxa such as amphibians and reptiles (apart from snakes) substantially delays ecological research in this region. Here, we compile an illustrated species list based on three years of research in and around the Lore Lindu National Park (LLNP) area and supplement it with data from the literature. In total, our survey and the literature review revealed 25 amphibian and 54 reptile species in five and 13 families, respectively. Our results highlight the LLNP area as an important herpetological endemism hotspot in the region. Appropriate utilization of species lists like this may facilitate capacity-building of local scientists and knowledgable local guides working in ecotourism.

Key to the Calamaria species of the Sulawesi region and the Moluccas (In part after Inger & Marx 1965, In den Bosch 1985, and de Lang & Vogel 2005) 1 Preocular absent......................................................................................................................................................... 9 - Preocular present......................................................................................................................................................... 2 2 Mental not touching anterior chin shields.................................................................................................................... 3 - Mental touching anterior chin shields........................................................................................................................... 5 3 Paraparietal surrounded by five scales and shields................................................................................ C. brongersmai - Paraparietal surrounded by six scales and shields........................................................................................................ 4 4 Eye diameter almost 1.5 times eye-mouth distance................................................................................. C. boesemani - Eye diameter slightly greater than eye-mouth distance.............................................................................. C. virgulata 5 First gular touching anterior chin shields ................................................................................................ C. acutirostris - First gular not touching anterior chin shields.............................................................................................................. 6 6 Paraparietal surrounded by five scales and shields ....................................................................................... C. muelleri - Paraparietal surrounded by six scales and shields....................................................................................................... 7 7 Tail thick, tapering abruptly at end ............................................................................................................. C. virgulata - Tail tapering gradually from base............................................................................................................................... 8 8 Ventrals yellow, immaculate except for dark lateral tips............................................................................. C. nuchalis - Ventrals dark, yellowish on posterior edges only .............................................................................................. C. curta 9 Mental touching anterior chin shields................................................................................................ C. apraeocularis - Mental not touching anterior chin shields.................................................................................................................. 10 10 Four supralabials..................................................................................................................................... C. longirostris - Five supralabials......................................................................................................................................................... 11 11 Pale collar absent ...................................................................................................................................... C. butonensis - Pale collar present....................................................................................................................................................... 12 12 ventrals 139 to 146 (males) or 148 to 165 (females), tail ratio in females 0.071 to 0.087, in males 0.125 to 0.150.... ................................................................................................................................................................. C. ceramensis - 157 (male) to 198 (female) ventrals, tail ratio in females 0.060, in males 0.103 ................................ C. banggaiensis

Key to the Calamaria species of the Sulawesi region and the Moluccas (In part after Inger & Marx 1965, In den Bosch 1985, and de Lang & Vogel 2005) 1 Preocular absent......................................................................................................................................................... 9 - Preocular present......................................................................................................................................................... 2 2 Mental not touching anterior chin shields.................................................................................................................... 3 - Mental touching anterior chin shields........................................................................................................................... 5 3 Paraparietal surrounded by five scales and shields................................................................................ C. brongersmai - Paraparietal surrounded by six scales and shields........................................................................................................ 4 4 Eye diameter almost 1.5 times eye-mouth distance................................................................................. C. boesemani - Eye diameter slightly greater than eye-mouth distance.............................................................................. C. virgulata 5 First gular touching anterior chin shields ................................................................................................ C. acutirostris - First gular not touching anterior chin shields.............................................................................................................. 6 6 Paraparietal surrounded by five scales and shields ....................................................................................... C. muelleri - Paraparietal surrounded by six scales and shields....................................................................................................... 7 7 Tail thick, tapering abruptly at end ............................................................................................................. C. virgulata - Tail tapering gradually from base............................................................................................................................... 8 8 Ventrals yellow, immaculate except for dark lateral tips............................................................................. C. nuchalis - Ventrals dark, yellowish on posterior edges only .............................................................................................. C. curta 9 Mental touching anterior chin shields................................................................................................ C. apraeocularis - Mental not touching anterior chin shields.................................................................................................................. 10 10 Four supralabials..................................................................................................................................... C. longirostris - Five supralabials......................................................................................................................................................... 11 11 Pale collar absent ...................................................................................................................................... C. butonensis - Pale collar present....................................................................................................................................................... 12 12 ventrals 139 to 146 (males) or 148 to 165 (females), tail ratio in females 0.071 to 0.087, in males 0.125 to 0.150.... ................................................................................................................................................................. C. ceramensis - 157 (male) to 198 (female) ventrals, tail ratio in females 0.060, in males 0.103 ................................ C. banggaiensis, Published as part of Koch, Andr��, Arida, Evy, Mcguire, Jimmy A., Iskandar, Djoko T. & B��hme, Wolfgang, 2009, A new species of Calamaria (Squamata: Colubridae) similar to C. ceramensis de Rooij, 1913, from the Banggai Islands, east of Sulawesi, Indonesia, pp. 19-30 in Zootaxa 2196 on pages 28-29, DOI: 10.5281/zenodo.189513, {"references":["Inger, R. F. & Marx, H. (1965) The systematics and evolution of the Oriental colubrid snakes of the genus Calamaria. Fieldiana Zoology, 49, 1 - 304.","In den Bosch, H. A. J. (1985) Snakes of Sulawesi: Checklist, key and additional biogeographical remarks. Zoologische Verhandelingen, 217, 3 - 50.","de Lang, R. & G. Vogel (2005) The Snakes of Sulawesi. A Field Guide to the Land Snakes of Sulawesi with Identification Keys. Edition Chimaira, 312 pp."]}

Cyrtodactylus nuaulu sp. nov. Figures 1���2 Holotype: MZB lace 2326 (F-num S 80) adult male, 26 /08/ 87, ~ 50m asl, Solea (2 �� 51 ' S, 129 �� 39 ' E), Manusela National Park, central Seram Island, Maluku Province, Indonesia, collected by P. Edgar and R. Lilley. Paratypes: MZB lace 2325 (F-num S 52) adult male, 20 / 10 / 87, MZB lace 2327 (F-num S 152) adult female, 13 /09/ 87 with same locality and collector details as holotype; MZB lace 2328 (F-num S 203) adult female 20 / 10 / 87 and MZB lace 2329 (F-num S 227) juvenile 29 / 10 / 87 from Waikawa, Manusela National Park, 3 km from Saunulu Village, inland from Japutih (3 �� 17 ' S 129 �� 31 ' E) southern Seram Island, Maluku Province, Indonesia. Diagnosis. Cyrtodactylus nuaulu sp. nov. can be distinguished from all other Melanesian and Wallacean Cyrtodactylus by the following unique combination of character states: moderate size (SVL up to 88.5mm); relatively slender body with robust head (HW/SVL 0.187���0.195); deep precloacal groove with low number of pores (6); femoral pores absent; subcaudal scales granular, not transversely enlarged; small dentate tubercles over the dorsum and along the lateral fold; whorls of prominent dentate tubercles extending to the tip of the tail; and dorsal colouration consisting of relatively few large indistinct transverse and/or longitudinal dark brown blotches. Description of holotype. A moderately large (81.7 SVL mm) and slender gecko. Head long (HL/SVL 0.253), moderately wide (HW/HL 0.744) and distinct from neck. Snout tapering to relatively blunt tip in dorsal profile, relatively long (longer than eye diameter), loreal region weakly inflated, interorbital region and top of snout strongly concave, canthus rostralis smoothly rounded. Eyes very large with vertical pupil. Supracillaries extending from anterior-ventral to posterior-dorsal edge of eye, longest at the anterior-dorsal corner. Ear opening small, dorso-ventrally flattened, and oriented at 45 degrees to apex of rictus with dorsal edge posteriormost, bordered by small indistinct ventral skinfold. Rostral approximately twice as wide as high, with short medial suture, widest at the ventral edge of the nares, bordered dorsally by right supranasal, larger rounded internasal, and damaged left supranasal. Nares bordered by first supralabial, rostral, first supranasal (point contact only) and series of three elongate postnasals. Supralabials: 10 on right lip and 11 on left, 7 or 8 to midpoint of eye, supralabials anterior to eye much broader than high and bordered dorsally by a single series of variably sized enlarged scales. Head scales small and granular, temporal and nuchal scales smaller than those on snout, scattered small conical tubercles on temporal and nuchal regions. Infralabials to rictus: 9 on right, 10 on left, all much broader than high, bordered by several rows of enlarged scales grading into small and granular gular scales. Mental triangular, approximately as wide as long, bordered by first infralabials and two diamond-shaped postmentals in contact for approximately 50 % of their length. Body elongate (TrL/SVL 0.494) with moderately distinct ventrolateral folds. Moderately tuberculate, lateral fold with distinct dentate tubercles separated from each other by 2���6 granules, posterior tubercles closer together. Dorsum with approximately fourteen rows (including lateral fold) of dentate tubercles. Dorsal scales small and granular. Ventral scales much larger than dorsal scales, increasing in size medially, arranged in approximately 51 rows at midpoint of body. Prominent raised straight precloacal groove surrounded by several rows of enlarged ventral scales, largest scales at base of groove containing 3 precloacal pores on each side. Distinctly enlarged rows of femoral scales absent. Forelimbs relatively elongate (FA/SVL 0.158), hindlimbs much more robust than forelimbs (CS/SVL 0.191). Lateral and dorsal surfaces of limbs with rows of dentate tubercles. Digits long and well developed, inflected at basal interphalageal joints; subdigital lamellae smooth, rounded, undivided and expanded proximal to joint inflection; large recurved claws sheathed by a dorsal and ventral scale. Slight basal webbing between digits II���IV on both manus and pes; lamellae on digits I /IV of manus 8 / 15 R 10 / 15 L and pes 12 / 17 R 9 / 15 L. Tail original, long and slender, tapering to point with distinct lateral groove extending approximately two thirds its length. Caudal scales granular, increasing in size ventrally, numerous rows of prominent dentate tubercles extending along all surfaces of tail for its full length, including 2 ventral rows. Hemipenal bulge swollen and prominent, left hemipenis everted, two enlarged postcloacal tubercles present at base of tail. Colouration. Dorsal ground colour light brownish grey with extensive fine brown flecking; a pair of almost continuous (broken just anterior to the hindlimbs) dark brown dorsolateral streaks extend from behind the eye to base of the tail; 2 darkish brown transverse blotches between the fore and hindlimbs. Nuchal region light brown with dark brown V-shaped blotch. Lateral regions light grey with small amounts of dark brown pigmentation forming indistinct scattered blotches and a faint discontinuous brown ventro-lateral stripe between fore and hindlimbs, strongest anteriorly and becoming very indistinct posteriorly. Venter grey with scattered light brown speckling, densest between the forelimbs. Head light grey dorsally, bordering relatively sharply against brown nuchal region, three small dark brown blotches forming Y-shape just posterior to orbital, additional dark brown longitudinal blotches medial to both eyes, above the rostrum and along the dorsal edge of the off-white supralabials. Supracillaries off-white and dark brown. Broad indistinct lateral band extends from orbital, above ear, and joins dorsolateral bands on the body. Infralabials off-white with scattered small dark brown blotches, ventral surface of lower jaw yellowishgrey with scattered indistinct greyish-brown flecking. Limbs dorsally light greyish brown with wide indistinct dark brown bands; bands on hindlimbs much lighter than forelimbs; digits mottled light and dark brown, with off-white bands proximal to claws. Ventrally limbs and digits greyish yellow with extensive brown speckling and blotching, especially on the crus. Tail with six wide dark brown bands and six (including tail tip) narrower light greyish to off-white bands; posteriormost dark bands slightly broken up by small amounts of light grey mottling. Variation. Comparative mensural and meristic data for the holotype and paratypes is given in Table 1. While broadly consistent, the colouration of the adult types shows significant differences (Fig. 2). All adult paratypes possess three dark to very dark brown transverse bands or blotches between the fore and hindlimbs (versus two in the holotype). The shape and definition of these marking also varies considerably; the holotype is the only specimen with dark dorsolateral stripes that extend to the legs, on other specimens these stripes tend to be lighter and at most extend to the midpoint of the body. The extent and darkness of other dorsal marking also varies, although all specimens possess a dark nuchal V- or Y-shape and at least some dark mottling or blotches between the eyes. The base dorsal colour of MZB lace 2328 is more brownish than grey. All types have at least some brown mottling on the ventral and lateral surfaces, however the density and extent of pigmentation again varies (the venter of MZB lace 2335 is particularly densely flecked). On the two adult individuals with original tails the number of bands varies from five to six (dark bands) to four to five (light bands); the tip of the original tail is dark in one of the paratypes (MZB lace 2325) as opposed to light in the holotype (MZB lace 2326). MZB lace 2327 has an almost complete original tail (1.5cm regrown at the tip) which was broken at collection, with five light and five dark bands. Approximately 60 % of the tail on paratype MZB lace 2328 is original with three off white bands and two greyish brown bands; the regrown section is yellowish off-white with extensive brown pigmentation, no tubercles and uniform scalation. MZB lace 2329 is a recently hatched juvenile, with original tail that was broken at collection and a circular patch of skin missing from around the left ear. The dorsal pattern on this specimen is simplified relative to the adult types and consists of two brown longitudinal dorsolateral streaks, a dark brown nuchal blotch and small brown blotch at the midpoint of the hindlegs. The lateral and ventral regions are also relatively plain with less brown pigmentation than the other types. Comparisons. The whorls of dentate tubercles extending to the tip of the original tail readily distinguish C. nuaulu sp. nov. from its two recognised Moluccan congeners; C. deveti and C. halmahericus. Cyrtodactylus deveti can be further distinguished by its more robust build, absence of a precloacal groove, possession of several rows of enlarged femoral scales, and much larger ventral caudal scales. Cyrtodactylus halmahericus, collected both sympatrically and on other Moluccan islands, are smaller (Cyrtodactylus ascribed to C. papuensis (Brongersma) from New Guinea and surrounding islands, lack enlarged tubercles on the tail and dorsum, are smaller (Cyrtodactylus by the presence of a precloacal groove. It can be further distinguished from the only other Melanesian species with enlarged tubercles along the tail, Cyrtodactylus serratus Kraus, (and related members of the C. loriae and C. louisiadensis species groups sensu R��sler et al. 2007) by its relatively small size and very small ventral caudal scales. Cyrtodactylus darmandvillei (Weber) from the Lesser Sundas lacks a precloacal groove, is much more robust and has much larger trihedral tubercles across the dorsum (see figure in De Rooji 1915). Cyrtodactylus wetariensis (Dunn), also from the lesser Sundas, is smaller Cyrtodactylus spinosus Linkem et al. from Sulawesi has long thin spines along the lateral fold, on the tail and on the postantefemoral region of the hindlimbs (Linkem et al. 2008). Other Indonesian Cyrtodactylus from islands to the west and north of Seram lack rings of enlarged tubercles extending the full length of the tail. Of the most geographically proximate species, C. marmoratus (Gray) (Java) and C. fumosus (M��ller) (Java and Sulawesi) can be distinguished by possessing a femoral pore series, while C. jellesmae (Boulenger) and C. wallacei Hayden et al. from Sulawesi lack a precloacal groove; all these species also have far more extensive and darker transverse banding or blotching across the dorsum (Hayden et al. 2008). Distribution and Natural History. Cyrtodactylus nuaulu sp. nov. is only known from two localities in lowland rainforest from the island of Seram. The forest around the type locality was dominated by Eucalyptus deglupta and Shorea sp. All adult types were collected by hand at night between 50���200cm above the ground, head down on trees and vines. The juvenile paratype was collected in a pit-trap at night. Other geckos collected in sympatry or near sympatry were Cyrtodactylus halmahericus, a probably undescribed small (Nactus sp. (listed as Cyrtodactylus pelagicus in Edgar & Lilley 1993) and Gehyra mutilata. Both collection localities are in Manusela National Park, which at the time of collection, was being threatened by illegal logging and conversion of forest to gardens. Eytmology. Named in honour of the Nuaulu people of south Seram. Cyrtodactylus nuaulu sp. nov. shows a combination of morphological characters that is very distinctive and does not strongly suggest a close relationship with any described Cyrtodactylus. While biogeographic studies and field surveys have found that the mammal, bird and frog fauna of Seram have a strong Melanesian influence (Edgar & Lilley 1993; Helgen 2003), most Melanesian Cyrtodactylus lack a precloacal groove, possess obvious femoral pores and have a relatively robust build. We tentatively suggest that Cyrtodactylus nuaulu sp. nov. is most likely to be allied to a number of smaller bodied, relatively gracile taxa with precloacal grooves distributed both to the west (C. cf marmoratus) and east (C. halmahericus, C. papuensis) of Wallace's Line. Nonetheless, we emphasise that the distinctive morphology of this species makes this association tenuous at best. Seram and surrounding islands (especially Buru) are recognised areas of endemism for birds (Stattersfield et al. 1998) and mammals (Flannery 1995; Helgen 2003). With the description of Cyrtodactylus nuaulu sp. nov. (known only from Seram) four reptile species are known to be endemic to this region of the Maluku Islands. The other endemic and near endemic species (also found on the neighbouring small island of Ambon) are the recently described monitor Varanus ceramboinensis (Philipp et al. 1999), Carlia leucotaenia and Calamaria ceramensis. Not surprisingly given the biogeographic position of Seram, these endemics include representatives of both Australasian (Carlia) and Asian (Calamaria) groups. As with mammals and birds, it seems that the herpetofauna of Seram and surrounding islands includes a small but significant and biogeographically interesting endemic component. It would also seem likely that further taxonomic and survey work in Seram and surrounding islands will add to this total. In particular the neighbouring large island of Buru remains very poorly explored and should be a priority for further surveys., Published as part of Oliver, Paul, Edgar, Paul, Iskandar, Djoko T. & Lilley, Ron, 2009, A new species of bent-toed gecko (Cyrtodactylus: Gekkonidae) from Seram Island, Indonesia, pp. 47-55 in Zootaxa 2115 on pages 48-54, DOI: 10.5281/zenodo.274887, {"references":["Rosler, H., Gunther, R. & Richards, S. J. (2007) Remarks on the morphology and taxonomy of geckos of the genus Cyrtodactylus Gray, 1827, occurring east of Wallacea, with description of two new species (Reptilia: Sauria: Gekkonidae). Salamandra, 43, 193 - 230.","Dunn, E. R. (1927) Results of the Douglas Burden Expedition to the island of Komodo. III. Lizards from the East Indies. American Museum Novitates, 288.","Linkem, C. W., McGuire, J. A., Hayden, C. J., Setiadi, M. I., Bickford, D. P. & Brown, R. M. (2008) A new species of benttoe gecko (Gekkonidae: Cyrtodactylus) from Sulawesi Island, eastern Indonesia. Herpetologica, 64, 224 - 234.","Edgar, P. & Lilley, R. (1993) Herpetofauna survey of Manusela National Park. Chapter 8. In: Edwards, I. D., McDonald, A. A. & Proctor, J. (Eds). Natural History of Seram. Intercept Ltd, Hampshire, England, pp. 131 - 141.","Helgen, K. M. (2003) A review of the rodent fauna of Seram, Moluccas, with the description of a new subspecies of mosaic-tailed rat, Melomys rufescens paveli. Journal of Zoology, London 261, 165 - 172.","Stattersfield, A. J., Corsby, M. J., Long, A. J. & Wege, D. C. (1998) Global directory of endemic bird areas. Cambridge UK: Birdlife International.","Philipp, K. M, Bohme, W. & Ziegler, T. (1999) The identity of Varanus indicus: redefinition and description of a sibling species co-existing at the type locality (Sauria: Varanidae: Varanus indicus group). Spixiana, 22, 273 - 287."]}

Oliver, Paul, Edgar, Paul, Iskandar, Djoko T., Lilley, Ron (2009): A new species of bent-toed gecko (Cyrtodactylus: Gekkonidae) from Seram Island, Indonesia. Zootaxa 2115: 47-55, DOI: 10.5281/zenodo.274887

Calamaria banggaiensis sp. nov. Figs. 2, 4, 6, and 8 Holotype. MZB Oph[idia]. 3230 (Field number AK0182), a female, collected by A. Koch and E. Arida at approximately 5 m elevation, southeast of Desa (= village) Banggai (1 �� 37 ��� 23 ������S, 123 �� 32 ��� 16 ������E), Pulau Banggai, Kepulauan Banggai, east of Central Sulawesi (Propinsi Sulawesi Tengah), Indonesia (Fig. 1), on 28 August 2005. Paratype. MZB Oph[idia]. 3755 (Field number JAM 8759) a male with everted hemipenes, collected by J. A. McGuire and D. T. Iskandar at 6 m above sea level, Kabupaten Bangkep (01�� 36 ��� 896 ������S, 123 �� 16 ��� 626 ������E), Pulau Peleng, Kepulauan Banggai, east of Central Sulawesi (Propinsi Sulawesi Tengah), Indonesia (Fig. 1), on 16 September 2007; body mass (in life) 3.0 g. Diagnosis. A new species of Calamaria that is distinguished from all other members of the genus by the following combination of characters (see below for specific comparisons): Preocular scales absent. Five supralabials (third and fourth contacting orbit) and five infralabials with the first pair meeting behind the mental so that the mental is separated from the anterior chin shields. Paraparietal surrounded by five scales and shields. Modified maxillary teeth. Smooth dorsal scales in 13 rows around the body. High number of ventral scales (157���198), a single anal plate, and 20���25 divided subcaudals. The dorsal colour pattern consists of pale brown with darker spots, a pale collar, and a lateral light and dark zigzag pattern along the body. A dark median streak on the underside of the tail is absent. Description of holotype and morphological variation. Features of the holotype are followed in parentheses by data from the paratype if different. The holotype is most probably a female due to the relatively small number of subcaudals and low tail length (as compared with C. ceramensis). Habitus vermiform. The head is blunt, not distinct from body. Snout-vent length 189 mm (199 mm), tail length 12 mm (23 mm). Ratio of tail length to total length 0.060 (0.103). The tail is short (moderately long in the male paratype), thick and slightly tapering to a point (tapered only at the end in male paratype). Seven modified maxillary teeth are visible. Portion of rostral scale visible from above; prefrontal shorter than frontal, touching first three supralabials; 13 dorsal rows of smooth scales; five supralabials, third and fourth entering orbit, fifth largest, second larger than first, third equal to first and fourth equal to second; frontal hexagonal, anterior portion compressed toward posterior and just longer than prefrontals, frontal about 2.5 times maximum width of supraocular; supraocular about equal to eye size; preocular absent; one postocular, taller than wide, as tall as eye diameter; eye diameter slightly larger than eye-mouth distance; loreal and supranasal absent; prefrontal shorter than frontal, touching the first, second, and third supralabials; nasal smaller than postocular; five infralabials, the first pair meet behind the mental preventing contact between mental and anterior chin shields; mental triangular; second pair of chin shields shorter than first, meeting in midline; paraparietals surrounded by five shields and scales; parietal about 1.8 times length of prefrontal; three gulars in midline between posterior chin shields and first ventral; first ventrals irregularly single or double, 198 (157) in total, strongly overlapping; anal plate single, broadened, with a small dark dot in the middle; 20 (25) divided subcaudals plus terminal spine. Colour in life: Body dorsally greyish-brown with irregularly scattered small dark spots all along the body and tail (ground colour darker in male paratype); head dark brown above with a weak pale, V-shaped collar bordered posteriorly by a dark brown band on the neck (collar not bordered by dark band in male paratype); parietals and paraparietals brown with pale markings; dark pigments covering upper one-third of supralabials and sutures between the shields, remainder of supralabials whitish; underside of head whitish with brown spots on the first three infralabials and the anterior chin shields; ventral side immaculate whitish-cream; laterally the pale ventral colouration reaches to the fourth dorsal scale in a zigzag pattern on the first half of the body; tail whitish-cream below, without a dark median streak. Hemipenes: The hemipenes are everted in the paratype, although they may be incompletely inflated. The hemipenes are simple relative to those of most snakes in that they are small (reaching only the fourth subcaudal), unicapitate with the sulcus spermaticus divided only near the terminus, and without elaborate ornamentation. Distally, each hemipenis has a pair of slightly enlarged lobes. Neither the lobes, nor the body of the organ has spines, although the basal two-thirds of each hemipenis is weakly plicate. There is a terminal knob-like projection between the termini of the sulcus spermaticus that might be capable of further eversion. Etymology. The specific epithet banggaiensis refers to the type locality, the Banggai Islands, east of Central Sulawesi. As vernacular names we suggest Banggai reed snake (English), Banggai Zwergschlange (German), and Calamaire de Banggai (French). Comparisons. The absence of a preocular scale is an important character that distinguishes Calamaria banggaiensis from the following 43 species in the genus: C. abstrusa Inger & Marx, 1965, C. acutirostris Boulenger, 1896, C. albiventer (Gray, 1834), C. battersbyi Inger & Marx, 1965, C. bicolor Dum��ril, Bibron & Dum��ril, 1854, C. bitorques Peters, 1872, C. boesemani Inger & Marx, 1965, C. borneensis Bleeker, 1860, C. brongersmai Inger & Marx, 1965, C. buchi Marx & Inger, 1955, C. crassa Lidth de Jeude, 1922, C. curta Boulenger, 1896, C. d��derleini Gough, 1902, C. eiselti Inger & Marx, 1965, C. everetti Boulenger, 1893, C. forcarti Inger & Marx, 1965, C. gervaisi Dum��ril & Bibron, 1854, C. gialaiensis Ziegler, Sang & Truong, 2008, C. grabowskyi Fischer, 1885, C. griswoldi Loveridge, 1938, C. hilleniusi Inger & Marx, 1965, C. ingeri Grismer, Kaiser & Yaakob, 2004, C. joloensis Taylor, 1922, C. lateralis Mocquard, 1890, C. lautensis de Rooij, 1917, C. leucogaster Bleeker, 1860, C. linnaei Boie, 1827, C. lumbricoidea Boie, 1827, C. lumholtzii Andersson, 1923, C. margaritophora Bleeker, 1860, C. melanota Jan, 1862, C. modesta Dum��ril & Bibron, 1854, C. muelleri Boulenger, 1896, C. nuchalis Boulenger, 1896, C. palavanensis Inger & Marx, 1965, C. pavimentata Dum��ril & Bibron, 1854, C. prakkei Lidth de Jeude, 1893, C. schlegeli Dum��ril, Bibron & Dum��ril, 1854, C. septentrionalis Boulenger, 1890, C. suluensis Taylor, 1922, C. sumatrana Edeling, 1870, and C. ulmeri Sackett, 1940, and C. virgulata Boie, 1827. Below we individually compare and distinguish the remaining species that share the absence of a preocular scale with C. banggaiensis. From the following non-Sulawesian or Moluccan Calamaria species that lack a preocular, C. banggaiensis differs from C. schmidti Marx & Inger, 1955, in having modified (vs. unmodified) maxillary teeth, five (vs. six) scales and shields around the paraparietal, and five (vs. four) supralabials. From C. javanica Boulenger, 1891, it differs in having the paraparietal surrounded by five scales (vs. six), having five (vs. four) supralabials, third and fourth (vs. second and third) entering the orbit. It differs from C. lovii Boulenger, 1887, and C. gracillima (G��nther, 1872) in that these species have either the second and third (C. lovii and C. gracillima) or only the third (only in C. lovii) supralabial entering the eye, whereas C. banggaiensis has the third and fourth supralabials in contact with the orbit. From C. alidae Boulenger, 1920, C. banggaiensis differs in having the paraparietal surrounded by five (vs. six) scales, and having seven (vs. nine to ten) modified maxillary teeth. In addition to the substantial geographical distance from Sumatra, the new species differs from C. mecheli Schenkel, 1901, in having a substantially different colour pattern lacking a dark vertebral stripe (as depicted by Inger & Marx 1965: 234), fewer ventral scales in males (157 vs. 174) and higher ratios of tail to total length in both sexes (0.103 vs. 0.116 in males and 0.060 vs. 0.046 to 0.049 in females, respectively). From the recently described C. thanhi Ziegler & Le, 2005, C. banggaiensis differs in having a light brown (vs. black in C. thanhi) background colour, five (vs. four) supralabials, and the third and fourth (vs. second and third in the Vietnamese species) supralabial touching the orbit. Among the Calamaria species lacking preocular scales, C. banggaiensis is most similar morphologically to the following species, all but one of which occur on Sulawesi, its satellite island of Buton, or in the adjacent Moluccan region. The new species is morphologically similar to C. ceramensis from the Moluccas, in the mental not contacting the anterior chin shields, the absence of preocular scales, prefrontal touching first three supralabials, five scales surrounding the paraparietals and the presence of a pale collar (Figs. 3, 5, 7, and 9). However, C. banggaiensis differs from C. ceramensis and the morphologically similar C. rebentischi Bleeker, 1860 from Borneo in having many more ventral scales (157 versus 139���146 in male and 198 vs. 148���165 in female C. ceramensis, and vs. 140 in male C. rebentischi), a smaller tail:total length ratio (0.060 vs. 0.071��� 0.087 in female and 0.103 vs. 0.125���0.15 in male C. ceramensis, and vs. 0.152 in the male type specimen of C. rebentischi), and by a larger eye diameter that slightly exceeds the eye-mouth distance (vs. equal to or smaller than eye-mouth distance in C. ceramensis [Fig. 7] and C. rebentischi, respectively). In addition, a dark median line on the ventral surface of the tail is sometimes present in C. ceramensis (see Fig. 5), but is not present on the holotype (Fig. 4) and paratype of C. banggaiensis. Calamaria banggaiensis is distinguished from C. longirostris Howard & Gillespie, 2007 (Buton Island), C. butonensis Howard & Gillespie, 2007 (Buton Island), and C. apraeocularis Smith, 1927 (southwestern peninsula of Sulawesi) by the lack of contact between the mental and the anterior pair of chin shields. The new taxon is further distinguished from C. apraeocularis by many more subcaudals in both sexes (20 vs. 10 in females and 25 vs. 18���19 in males), by having five (vs. six) shields and scales surrounding the paraparietal, by significantly longer tails in both sexes (0.060 vs. 0.030���0.035 in females and 0.103 vs. 0.067���0.068 in males), and by having a pale collar. Moreover, C. banggaiensis differs further from C. longirostris in having five (vs. four) supralabials, and from C. butonensis in having more ventrals (198 vs. 141���177) and subcaudals (20 vs. 14���18) in females. Furthermore, the two Calamaria species from Buton Island show a different colour pattern in that they lack a pale collar. Distribution, habitat, and natural history. Currently, Calamaria banggaiensis is known only from the islands of Banggai and Peleng east of Central Sulawesi (Fig. 1). It is reasonable to expect this species to occur on smaller nearby islands such as Labobo and Bangkurung that are merely separated by shallow sea ways. Occurrence of C. banggaiensis on Sulawesi, however, is questionable due to the deep channel that separates the Banggai Archipelago from Sulawesi precluding a land connection during glacial maxima, and the associated faunistic differences in the assemblage of amphibians and reptiles of Sulawesi mainland and the Banggai island group (unpublished data). The type specimens were both collected in the afternoon under surface objects (the holotype was collected under a decomposing log, the paratype under a rock). In the former case, the specimen was found in a cacaoplantation near a river (Banggai Island), the second at a site that was characterized by a combination of large trees and cacao understory (Peleng Island). Both specimens were discovered only a few meters above sea level. As for most Calamaria species, nearly nothing is known about the biology of this newly described taxon and further investigations are needed. In concordance with the secretive and borrowing lifestyle of Calamaria, the Peleng specimen regurgitated an earthworm after capture. Other amphibians and reptiles that were found in the anthropogenically-influenced habitat of C. banggaiensis on Banggai Island are Kaloula cf. baleata, Platymantis papuensis, Draco rhytisma, Emoia caeruleocauda, Lamprolepis smaragdina ssp., Lipinia cf. infralineolata, and Varanus salvator ssp. (Koch et al. 2007 a)., Published as part of Koch, Andr��, Arida, Evy, Mcguire, Jimmy A., Iskandar, Djoko T. & B��hme, Wolfgang, 2009, A new species of Calamaria (Squamata: Colubridae) similar to C. ceramensis de Rooij, 1913, from the Banggai Islands, east of Sulawesi, Indonesia, pp. 19-30 in Zootaxa 2196 on pages 21-27, DOI: 10.5281/zenodo.189513, {"references":["Inger, R. F. & Marx, H. (1965) The systematics and evolution of the Oriental colubrid snakes of the genus Calamaria. Fieldiana Zoology, 49, 1 - 304.","Ziegler, T., Sang, N. V. & Truong, N. Q. (2008) A new Reed Snake of the Genus Calamaria Boie (Squamata: Colubridae) from Vietnam. Current Herpetology, 27, 71 - 80.","Grismer, L. L., Kaiser, H. & Yaakob, N. S. (2004) A new species of reed snake of the genus Calamaria H. Boie, 1827, from Pulau Tioman, Pahang, West Malaysia. Hamadryad, 28, 1 - 6.","Boie, F. (1827) Bemerkungen uber Merrem's Versuch eines Systems der Amphibien. 1820. Marburg. Erste Lieferung: Ophidier. Isis von Oken, 20 (10), 508 - 566.","Ziegler, T. & Le, K. Q. (2005) A new species of reed snake, Calamaria (Squamata: Colubridae), from the Central Truong Son (Annamite mountain range), Vietnam. Zootaxa, 1042, 27 - 38.","Howard, S. D. & Gillespie, G. (2007) Two New Calamaria (Serpentes) Species from Sulawesi, Indonesia. Journal of Herpetology, 41 (2), 237 - 242.","Koch, A., Arida, E. & Bohme, W. (2007 a) Zwischenbericht uber die Herpetofauna Sulawesis unter besonderer Berucksichtigung der Gattung Varanus: phylogeographische Beziehungen zu angrenzenden Gebieten. Elaphe, 15 (3), 42 - 52."]}

Calamaria banggaiensis sp. nov. Figs. 2, 4, 6, and 8 Holotype. MZB Oph[idia]. 3230 (Field number AK0182), a female, collected by A. Koch and E. Arida at approximately 5 m elevation, southeast of Desa (= village) Banggai (1 ° 37 ’ 23 ’’S, 123 ° 32 ’ 16 ’’E), Pulau Banggai, Kepulauan Banggai, east of Central Sulawesi (Propinsi Sulawesi Tengah), Indonesia (Fig. 1), on 28 August 2005. Paratype. MZB Oph[idia]. 3755 (Field number JAM 8759) a male with everted hemipenes, collected by J. A. McGuire and D. T. Iskandar at 6 m above sea level, Kabupaten Bangkep (01° 36 ’ 896 ’’S, 123 ° 16 ’ 626 ’’E), Pulau Peleng, Kepulauan Banggai, east of Central Sulawesi (Propinsi Sulawesi Tengah), Indonesia (Fig. 1), on 16 September 2007; body mass (in life) 3.0 g. Diagnosis. A new species of Calamaria that is distinguished from all other members of the genus by the following combination of characters (see below for specific comparisons): Preocular scales absent. Five supralabials (third and fourth contacting orbit) and five infralabials with the first pair meeting behind the mental so that the mental is separated from the anterior chin shields. Paraparietal surrounded by five scales and shields. Modified maxillary teeth. Smooth dorsal scales in 13 rows around the body. High number of ventral scales (157–198), a single anal plate, and 20–25 divided subcaudals. The dorsal colour pattern consists of pale brown with darker spots, a pale collar, and a lateral light and dark zigzag pattern along the body. A dark median streak on the underside of the tail is absent. Description of holotype and morphological variation. Features of the holotype are followed in parentheses by data from the paratype if different. The holotype is most probably a female due to the relatively small number of subcaudals and low tail length (as compared with C. ceramensis). Habitus vermiform. The head is blunt, not distinct from body. Snout-vent length 189 mm (199 mm), tail length 12 mm (23 mm). Ratio of tail length to total length 0.060 (0.103). The tail is short (moderately long in the male paratype), thick and slightly tapering to a point (tapered only at the end in male paratype). Seven modified maxillary teeth are visible. Portion of rostral scale visible from above; prefrontal shorter than frontal, touching first three supralabials; 13 dorsal rows of smooth scales; five supralabials, third and fourth entering orbit, fifth largest, second larger than first, third equal to first and fourth equal to second; frontal hexagonal, anterior portion compressed toward posterior and just longer than prefrontals, frontal about 2.5 times maximum width of supraocular; supraocular about equal to eye size; preocular absent; one postocular, taller than wide, as tall as eye diameter; eye diameter slightly larger than eye-mouth distance; loreal and supranasal absent; prefrontal shorter than frontal, touching the first, second, and third supralabials; nasal smaller than postocular; five infralabials, the first pair meet behind the mental preventing contact between mental and anterior chin shields; mental triangular; second pair of chin shields shorter than first, meeting in midline; paraparietals surrounded by five shields and scales; parietal about 1.8 times length of prefrontal; three gulars in midline between posterior chin shields and first ventral; first ventrals irregularly single or double, 198 (157) in total, strongly overlapping; anal plate single, broadened, with a small dark dot in the middle; 20 (25) divided subcaudals plus terminal spine. Colour in life: Body dorsally greyish-brown with irregularly scattered small dark spots all along the body and tail (ground colour darker in male paratype); head dark brown above with a weak pale, V-shaped collar bordered posteriorly by a dark brown band on the neck (collar not bordered by dark band in male paratype); parietals and paraparietals brown with pale markings; dark pigments covering upper one-third of supralabials and sutures between the shields, remainder of supralabials whitish; underside of head whitish with brown spots on the first three infralabials and the anterior chin shields; ventral side immaculate whitish-cream; laterally the pale ventral colouration reaches to the fourth dorsal scale in a zigzag pattern on the first half of the body; tail whitish-cream below, without a dark median streak. Hemipenes: The hemipenes are everted in the paratype, although they may be incompletely inflated. The hemipenes are simple relative to those of most snakes in that they are small (reaching only the fourth subcaudal), unicapitate with the sulcus spermaticus divided only near the terminus, and without elaborate ornamentation. Distally, each hemipenis has a pair of slightly enlarged lobes. Neither the lobes, nor the body of the organ has spines, although the basal two-thirds of each hemipenis is weakly plicate. There is a terminal knob-like projection between the termini of the sulcus spermaticus that might be capable of further eversion. Etymology. The specific epithet banggaiensis refers to the type locality, the Banggai Islands, east of Central Sulawesi. As vernacular names we suggest Banggai reed snake (English), Banggai Zwergschlange (German), and Calamaire de Banggai (French). Comparisons. The absence of a preocular scale is an important character that distinguishes Calamaria banggaiensis from the following 43 species in the genus: C. abstrusa Inger & Marx, 1965, C. acutirostris Boulenger, 1896, C. albiventer (Gray, 1834), C. battersbyi Inger & Marx, 1965, C. bicolor Duméril, Bibron & Duméril, 1854, C. bitorques Peters, 1872, C. boesemani Inger & Marx, 1965, C. borneensis Bleeker, 1860, C. brongersmai Inger & Marx, 1965, C. buchi Marx & Inger, 1955, C. crassa Lidth de Jeude, 1922, C. curta Boulenger, 1896, C. döderleini Gough, 1902, C. eiselti Inger & Marx, 1965, C. everetti Boulenger, 1893, C. forcarti Inger & Marx, 1965, C. gervaisi Duméril & Bibron, 1854, C. gialaiensis Ziegler, Sang & Truong, 2008, C. grabowskyi Fischer, 1885, C. griswoldi Loveridge, 1938, C. hilleniusi Inger & Marx, 1965, C. ingeri Grismer, Kaiser & Yaakob, 2004, C. joloensis Taylor, 1922, C. lateralis Mocquard, 1890, C. lautensis de Rooij, 1917, C. leucogaster Bleeker, 1860, C. linnaei Boie, 1827, C. lumbricoidea Boie, 1827, C. lumholtzii Andersson, 1923, C. margaritophora Bleeker, 1860, C. melanota Jan, 1862, C. modesta Duméril & Bibron, 1854, C. muelleri Boulenger, 1896, C. nuchalis Boulenger, 1896, C. palavanensis Inger & Marx, 1965, C. pavimentata Duméril & Bibron, 1854, C. prakkei Lidth de Jeude, 1893, C. schlegeli Duméril, Bibron & Duméril, 1854, C. septentrionalis Boulenger, 1890, C. suluensis Taylor, 1922, C. sumatrana Edeling, 1870, and C. ulmeri Sackett, 1940, and C. virgulata Boie, 1827. Below we individually compare and distinguish the remaining species that share the absence of a preocular scale with C. banggaiensis. From the following non-Sulawesian or Moluccan Calamaria species that lack a preocular, C. banggaiensis differs from C. schmidti Marx & Inger, 1955, in having modified (vs. unmodified) maxillary teeth, five (vs. six) scales and shields around the paraparietal, and five (vs. four) supralabials. From C. javanica Boulenger, 1891, it differs in having the paraparietal surrounded by five scales (vs. six), having five (vs. four) supralabials, third and fourth (vs. second and third) entering the orbit. It differs from C. lovii Boulenger, 1887, and C. gracillima (Günther, 1872) in that these species have either the second and third (C. lovii and C. gracillima) or only the third (only in C. lovii) supralabial entering the eye, whereas C. banggaiensis has the third and fourth supralabials in contact with the orbit. From C. alidae Boulenger, 1920, C. banggaiensis differs in having the paraparietal surrounded by five (vs. six) scales, and having seven (vs. nine to ten) modified maxillary teeth. In addition to the substantial geographical distance from Sumatra, the new species differs from C. mecheli Schenkel, 1901, in having a substantially different colour pattern lacking a dark vertebral stripe (as depicted by Inger & Marx 1965: 234), fewer ventral scales in males (157 vs. 174) and higher ratios of tail to total length in both sexes (0.103 vs. 0.116 in males and 0.060 vs. 0.046 to 0.049 in females, respectively). From the recently described C. thanhi Ziegler & Le, 2005, C. banggaiensis differs in having a light brown (vs. black in C. thanhi) background colour, five (vs. four) supralabials, and the third and fourth (vs. second and third in the Vietnamese species) supralabial touching the orbit. Among the Calamaria species lacking preocular scales, C. banggaiensis is most similar morphologically to the following species, all but one of which occur on Sulawesi, its satellite island of Buton, or in the adjacent Moluccan region. The new species is morphologically similar to C. ceramensis from the Moluccas, in the mental not contacting the anterior chin shields, the absence of preocular scales, prefrontal touching first three supralabials, five scales surrounding the paraparietals and the presence of a pale collar (Figs. 3, 5, 7, and 9). However, C. banggaiensis differs from C. ceramensis and the morphologically similar C. rebentischi Bleeker, 1860 from Borneo in having many more ventral scales (157 versus 139–146 in male and 198 vs. 148–165 in female C. ceramensis, and vs. 140 in male C. rebentischi), a smaller tail:total length ratio (0.060 vs. 0.071– 0.087 in female and 0.103 vs. 0.125–0.15 in male C. ceramensis, and vs. 0.152 in the male type specimen of C. rebentischi), and by a larger eye diameter that slightly exceeds the eye-mouth distance (vs. equal to or smaller than eye-mouth distance in C. ceramensis [Fig. 7] and C. rebentischi, respectively). In addition, a dark median line on the ventral surface of the tail is sometimes present in C. ceramensis (see Fig. 5), but is not present on the holotype (Fig. 4) and paratype of C. banggaiensis. Calamaria banggaiensis is distinguished from C. longirostris Howard & Gillespie, 2007 (Buton Island), C. butonensis Howard & Gillespie, 2007 (Buton Island), and C. apraeocularis Smith, 1927 (southwestern peninsula of Sulawesi) by the lack of contact between the mental and the anterior pair of chin shields. The new taxon is further distinguished from C. apraeocularis by many more subcaudals in both sexes (20 vs. 10 in females and 25 vs. 18–19 in males), by having five (vs. six) shields and scales surrounding the paraparietal, by significantly longer tails in both sexes (0.060 vs. 0.030–0.035 in females and 0.103 vs. 0.067–0.068 in males), and by having a pale collar. Moreover, C. banggaiensis differs further from C. longirostris in having five (vs. four) supralabials, and from C. butonensis in having more ventrals (198 vs. 141–177) and subcaudals (20 vs. 14–18) in females. Furthermore, the two Calamaria species from Buton Island show a different colour pattern in that they lack a pale collar. Distribution, habitat, and natural history. Currently, Calamaria banggaiensis is known only from the islands of Banggai and Peleng east of Central Sulawesi (Fig. 1). It is reasonable to expect this species to occur on smaller nearby islands such as Labobo and Bangkurung that are merely separated by shallow sea ways. Occurrence of C. banggaiensis on Sulawesi, however, is questionable due to the deep channel that separates the Banggai Archipelago from Sulawesi precluding a land connection during glacial maxima, and the associated faunistic differences in the assemblage of amphibians and reptiles of Sulawesi mainland and the Banggai island group (unpublished data). The type specimens were both collected in the afternoon under surface objects (the holotype was collected under a decomposing log, the paratype under a rock). In the former case, the specimen was found in a cacaoplantation near a river (Banggai Island), the second at a site that was characterized by a combination of large trees and cacao understory (Peleng Island). Both specimens were discovered only a few meters above sea level. As for most Calamaria species, nearly nothing is known about the biology of this newly described taxon and further investigations are needed. In concordance with the secretive and borrowing lifestyle of Calamaria, the Peleng specimen regurgitated an earthworm after capture. Other amphibians and reptiles that were found in the anthropogenically-influenced habitat of C. banggaiensis on Banggai Island are Kaloula cf. baleata, Platymantis papuensis, Draco rhytisma, Emoia caeruleocauda, Lamprolepis smaragdina ssp., Lipinia cf. infralineolata, and Varanus salvator ssp. (Koch et al. 2007 a).

RANA ESCHATIA SP. NOV. (Previously referred to as Thailand morphotype) Rana labialis Smith, 1916: 168. Rana chalconota Smith, 1930: 109. Holotype THNHM 05677 (field number 66721), an adult female from Ngao Falls National Park (9°56′N / 98°43′E), Ranong Province, Thailand. Collected on a gravel bank 0.1 m from the edge of a stream in secondary forest, 26.xi.2004, by Jennifer Sheridan and Tanya Chan-ard. Paratypes From the type locality FMNH 268523, 268526-28, 268530 (adult males with nuptial pads) FMNH 268524, 268529 (adult females with convoluted oviducts), FMNH 268521, 268525 (juveniles). Etymology Specific name from eschatia, Gr., outskirt, referring to distribution at the edge of the geographical range of the group. Referred material Thailand: FMNH 268852-54, 268856-57 Khao Luang National Park (8°30′N / 99°45′E), Nakhon Si Thammarat Prov.; FMNH 268858, 268860, 268869 Khao Phanom Bencha National Park (8°14′N / 99°E), Krabi Prov.; FMNH 268872, 268874-84 Khao Sok National Park (8°56′N / 98°34′E), Surat Thani Prov.; FMNH 268531-4, 268536-9, THNHM 05690, 05695 Kaeng Krung National Park (9°34′N / 98°49′E), Surat Thani Prov.. Diagnosis A moderate-sized species of the chalconota group with males up to 40 mm SVL, females up to 57 mm, no dorsal spotting, relatively wide head (HW / SVL usually> 0.305), relatively long leg (T / SVL usually> 0.575) and males with constricted or divided nuptial pads. Description Habitus slender, head slightly wider than trunk, legs long. Head triangular; snout obtusely pointed, rounded in profile, projecting beyond lower jaw, longer than diameter of eye; nostril lateral, very close to tip of snout; canthus angular, not constricted; lores concave, vertical; interobital wider than upper eyelid and internarial; tympanum distinct, about two-thirds eye diameter in females, slightly larger in males, inside its rim the tympanum is slightly depressed relative to the surface of the temporal region; vomerine teeth in oblique groups, gap between groups less than length of one group and equal to distance from choana. Fingers long, third finger longer than snout; fingers without webbing; second and third fingers with narrow, movable fold of skin along medial margins; tips of three outer fingers with wide discs, that of third finger almost equal diameter of tympanum in female; disc of first finger much narrow than that of second; all discs with circummarginal grooves; subarticular tubercles conspicuous; third finger with two small supernumerary tubercles, bases of second and fourth fingers with a single supernumerary tubercle. Tips of toes expanded into discs smaller than those of outer fingers, but with circummarginal grooves; webbing extensive, reaching discs of first three toes on lateral margins and disc of fifth toe medially; fourth toe webbed to distal subarticular tubercle medially and slightly beyond that laterally; no dermal ridge along outer margins of first and fifth toes; a low oval inner metatarsal tubercle and a round outer one. Skin of back granular, in males granules weakly spinose; dorsolateral fold distinct, low; ventral surfaces smooth, except weakly rugose at rear of abdomen; rictal glands present. Colour in preservative dark brown dorsally and laterally; no black spots on dorsal surfaces; ventral surfaces cream-coloured or white; in some individuals throat with round dark spots; limbs without dark crossbars; rear of thigh dark brown with indistinct lighter round areas. Measurements (mm) of holotype: SVL 55.6, T 31.0, HW 16.0, HL 19.8, TYM 4.6, DF3 3.2. Variation Adult females 42.8–56.6 mm, mean 47.57 ± 0.96 mm (N = 17), males 30.6–39.6 mm, mean 34.14 ± 0.39 mm (N = 22). Variation in body proportions given in Table 8. Relative tympanum diameter in females 0.077 –0.095, in males 0.096 –0.127. The sexes do not differ in relative head width; HW/ SVL in females 0.288 –0.325 (median 0.308), in males 0.275 –0.326 (median 0.310). All males have constricted or divided nuptial pads and vocal sac openings at the corners of the mouth. Comparisons Males of Rana eschatia are most similar in size to those of R. labialis (Selangor) and R. parvaccola (Table 2), but females of eschatia are larger than females of those two (Tables 2 and 6). Rana eschatia also differs from those two species in the absence of black spots on the back and in having a wider head in both sexes (Tables 2 and 6). Rana eschatia is larger than R. raniceps (both sexes) and has a longer leg (T / SVL) and a smaller tympanum in males (Tables 2 and 6). Rana eschatia is smaller than R. megalonesa, R. rufipes and R. chalconota and differs from the latter two in the length of the tibia (T / SVL) and in the form of the nuptial pad, which is constricted or divided only in eschatia.

RANA RANICEPS(PETERS, 1871) (Previously referred to as Borneo Small morphotype) Polypedates raniceps Peters, 1871: 580 – Sarawak. Rana raniceps Iskandar & Colijn, 2000: 91; Stuart et al., 2006: 473. Rana (Chalcorana) raniceps Dubois, 1992: 328. Rana chalconota raniceps Inger, 1966: 177. Material examined MSNG 29376 lectotype (see Capocaccia, 1957) from ‘ Sarawak;’ Bintulu Division: Labang Forest Reserve (3°21′N / 113°27′E) FMNH 148083-219; Samarakan (2°56′N / 113°07′E) FMNH 267965-66; Bukit Sarang (2°39′N / 113°03′E) FMNH 267958-64. The paralectotype designated by Capocaccia (MSNG 50536) is a rhacophorid, with intercalary cartilages, a distinct projection at the heel and the entire abdomen coarsely granular, but no outer metatarsal tubercle. Diagnosis SVL of lectotype female 40.4 mm, other females 33.1–42.3 mm, mean 38.20 ± 0.48 mm (N = 30); males 27.6–34.1 mm, mean 30.35 ± 0.93 mm (N = 8). DF3 / SVL 0.055 –0.068, median 0.060. Back usually without dark spots. Males with nuptial pad constricted; male TYM/ SVL 0.089 –0.127 (N = 6). Descriptive notes The head is triangular and the snout slightly projecting. Inside its rim, the tympanum is slightly depressed relative to the surface of the temporal region. The pineal body is faintly visible and is in line with the anterior borders of the upper eyelids. The skin of the back is granular and in males is set with many fine spinules. The hind limb is without crossbars in most preserved specimens. Comparisons This is the smallest member of the chalconota group, differing significantly (P ³ 0.001, both sexes) from all other species except males of parvaccola (see below) (Tables 2 and 6). It differs from the two species from Padang, West Sumatra (parvaccola and rufipes see below), labialis and co-occurring R. megalonesa (see below) samples in the low frequency of dark dorsal spots. The uncorrected pairwise sequence divergence between R. raniceps and the co-occurring R. megalonesa (see below) is 13.11–13.97% (Table 4). This species is known only from low-lying areas of west-central Sarawak., Published as part of Inger, Robert F., Stuart, Bryan L. & Iskandar, Djoko T., 2009, Systematics of a widespread Southeast Asian frog, Rana chalconota (Amphibia: Anura: Ranidae), pp. 123-147 in Zoological Journal of the Linnean Society 155 (1) on pages 135-136, DOI: 10.1111/j.1096-3642.2008.00440.x, http://zenodo.org/record/5445984, {"references":["Peters W. 1871. Uber neue Reptilien aus Ostafrica und Sarawak (Borneo), vorzuglich aus der Sammlung des Hrn. Marquis J. Doria zu Genua. Monatsbericht der Koniglichten Akademie der Wissenschaften zu Berlin 1871: 566 - 581.","Iskandar DT, Colijn E. 2000. Preliminary checklist of Southeast Asia and New Guinean Herpetfauna [sic]. I. Amphibians. Truebia 31 (Suppl.): 1 - 133.","Stuart BL, Inger RF, Voris HK. 2006. High level of cryptic species diversity revealed by sympatric lineages of Southeast Asian forest frogs. Biology Letters 2: 470 - 474.","Dubois A. 1992. Notes sur la classification des Ranidae (Amphibiens Anoures). Bulletin Mensuel de la Societe Linneenne de Lyon 61: 305 - 352.","Inger RF. 1966. The systematics and zoogeography of the Amphibia of Borneo. Fieldiana: Zoology 52: 1 - 402.","Capocaccia L. 1957. Catalogo dei tipi di Amfibi del Museo Civico di Storia Naturale di Genova. Annali del Museo Civico di Storia Naturale di Genova 59: 208 - 222."]}

RANA MEGALONESA SP. NOV. (Previously referred to as Borneo Large morphotype) Rana chalconota (part) Boulenger, 1920: 201; van Kampen, 1923: 217. Rana chalconota raniceps Inger, 1966: 177. Rana cf. chalconota Stuart et al., 2006: 473. Holotype FMNH 267821, an adult female from Bukit Sarang (2°39′N / 113°03′E), Bintulu Division, Sarawak (Borneo), Malaysia. Collected in a freshwater swamp forest (20 m a.s.l.) 1 m above ground on a shrub, 11.xi.2004, by Freddy Paulus and Patrick Francis. Paratypes From type locality FMNH 267814-15, 267818, 267825 adult males with nuptial pads, FMNH 267816, 267819, 267824 adult females with convoluted, enlarged oviducts; FMNH 267817, 267820, 267822- 23 subadult females. Etymology Specific name from megalo-, Gr., large, and nesos, Gr., island, referring to its distribution on the large island of Borneo. Referred material Sarawak: Belaga District, Sg. Segaham (2°44′N / 113°55′E) FMNH 220474, 220477-78, 220484, 220492, 220512-13, 220526, 220541-43, 220547, 220549, 220551-52, 220554; Kapit District, Nanga Tekalit (1°37′N / 113°35′E) FMNH 220264, 220267-68, 220286, 220297, 220379, 220381, 220383, 220396, 220399, 220404, 220417, 220434, 220447, 220559-60, 220563- 64, 220568, 220570, 220572-74, 220576-77, 220579, 220581, 220586, 222955-56; Bintulu Division, Labang Forest Reserve (3°21′N / 113°27′E) FMNH 148203-07; Bintulu Division, Sg. Pesu camp (3°07′N / 113°48′E) FMNH 156607, 156610-11, 156622, 156627, 156631, 156634-35, 156638, 156640, 156643, 156653-56, 156658, 156660, 156666, 156668, 156674-77, 156680- 84, 156687, 156709-13, 156716, 156718, 156720, 156729-31, 156735, 156741, 156747, 156749, 156756- 59, 156762, 156765-68. Sabah: Kota Marudu District, Marak Parak (6°18′N / 116°42′E) FMNH 235639-45; Lahad Datu District, Danum Valley Research Centre (5°12′N / 117°50′E) FMNH 203953-62, 203965, 203969- 71, 203974-78, 203980, 203983, 203985, 203987-88, 203991-92; Sipitang District, Mendolong (4°54′N / 115°42′E) FMNH 128334, 238336, 238348, 238362, 242797-98, 242801, 242804, 242806-07, 242811; Tawau District, Bukit Tawau Park (4°37′N / 117°54′E) FMNH 248339-42, 248345-46, 248348-49; Tawau District, Kalabakan (4°25′N / 117°30′E) FMNH 76694, 76696, 76702, 76705, 76715-16, 76718, 76722, 76733, 76738, 76742-43, 76753, 76762-64, 76770, 76779, 76781-82, 76784-86, 76789. Diagnosis A large-sized member of the R. chalconota group; distinguished from other forms by combination of females usually> 50 mm, males> 35 mm; T / SVL usually> 0.56, HW / SVL usually> 0.30, DF3/ SVL> 0.06; males with nuptial pad constricted or divided and with weak humeral gland discernible only upon dissection. Description Habitus moderately slender, head slightly wider than trunk, legs long. Head triangular; snout obtusely pointed, projecting beyond lower jaw, longer than diameter of eye; nostril on side of snout, closer to tip of snout than to eye; canthus angular, not constricted; lores concave, weakly sloping; interorbital wider than upper eyelid and wider than internarial; pineal body faintly visible, between anterior corners of upper eyelids; tympanum distinct, about two-thirds eye diameter in female, slightly larger in males, slightly depressed relative to surface of temporal region; vomerine teeth in short, oblique groups between choanae, distance between groups shorter than distance from choanae. Fingers long, length of third finger equal to distance from rear of eye to nostril; fingers without webbing; second and third fingers with narrow, movable fold of skin on medial margins; tips of three outer fingers with wide discs, that of third finger about two-thirds diameter of tympanum in female, disc of first finger about half width of disc of second finger, all discs with circummarginal grooves; subarticular tubercles conspicuous, rounded; base of third finger with one or two supernumerary tubercles, bases of second and fourth fingers with one supernumerary tubercle. Tips of toes expanded into discs smaller than those of fingers, but with circummarginal grooves; webbing extensive, to base of discs on lateral margins of three inner toes and on medial margin of fifth, to base of disc on medial margin of fourth toe or between disc and distal subarticular tubercle; a narrow ridge of skin medially along first toe and a similar one along outer edge of last joint of fifth toe; a low, oval inner metatarsal tubercle and a distinct, round outer one. Skin of back weakly granular with scattered colourless spinules in females; males with densely crowded, taller spinules on all dorsal surfaces including head and eyelid, similar spinules on lores; a distinct, but low dorsolateral fold; ventral surface of body smooth, weakly rugose at rear of abdomen; a ridge-like rictal glandular swelling followed after a narrow gap by a glandular swelling above the axilla. Colour in preservative brown above and on sides, darker on side of head, upper lip white; many scattered dark spots on back and usually on head; ventral surfaces white, throat and chest with or without small dark spots; hind limb without dark crossbars in most preserved individuals; rear of thigh brown with faint, round lighter markings. Measurements (mm) of holotype: SVL 53.8, T 28.4, HW 16.6, HL 21.1, TYM 4.4, DF3 3.4. Variation Females 45.4–65.6 mm, mean 53.66 ± 0.45 mm (N = 115); males 33.3–48.2 mm, mean 39.21 ± 0.34 mm (N = 113). DF3/ SVL 0.054 –0.076, median 0.064 (N = 104). TYM/ SVL of males 0.089 –0.135, median 0.112 (N = 76). Humeral gland in males usually detectable only by dissection. Frequency of dark spotting on back varies among samples. In two samples from eastern Sabah dorsal spots were present in 18 of 28 frogs; in two samples from western Sabah dorsal spots were present in only seven of 29. The spots were present in two-thirds of frogs from the Bintulu Division of westcentral Sarawak but in only four of 90 from southeastern Sarawak. Frequency of constriction of the nuptial pad of males also varies. In frogs from eastern Sabah (three localities) the nuptial pad was constricted or divided in 22 of 28 males. The frequency of constriction in males from western Sabah (two localities) was five of 13. In males from south-eastern Sarawak (three localities) the frequency of constricted or divided nuptial pads was 24 of 51 individuals. The nuptial pad was constricted or divided in four of eight males from the Bintulu Division, west-central Sarawak. Comparisons The difference between this species and the cooccurring R. raniceps in size is striking. The mean SVL of males of R. megalonesa is roughly 10 mm larger than that of R. raniceps and the difference between means of females is almost 15 mm (Table 2). Individuals of Rana megalonesa that co-occur with R. raniceps differ from the latter in higher frequency of dark spots on the back. Although it is a large form of the chalconota group, females of R. megalonesa are smaller than those of both R. rufipes (see below) and Javan R. chalconota and its males smaller than those of R. rufipes (Tables 2 and 6). In addition to the size difference, the new species also differs from R. rufipes in having a relatively larger tympanum (Tables 2 and 6) and in the form of the nuptial pad (not constricted or divided in R. rufipes). Both males and females of R. megalonesa are larger than those of R. labialis, R. eschatia and R. parvaccola (see below). Relative head width (HW / SVL) of R. megalonesa is larger than that of R. labialis and R. parvaccola. Relative width of the tympanum (TYM / SVL) of R. megalonesa is larger than that of R. labialis in both sexes (Tables 2 and 6). The uncorrected pairwise sequence divergence between R. megalonesa and the co-occurring R. raniceps is 13.11–13.97% (Table 4).

Inger, Robert F., Stuart, Bryan L., Iskandar, Djoko T. (2009): Systematics of a widespread Southeast Asian frog, Rana chalconota (Amphibia: Anura: Ranidae). Zoological Journal of the Linnean Society 155 (1): 123-147, DOI: 10.1111/j.1096-3642.2008.00440.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2008.00440.x

RANA RUFIPES SP. NOV. (Previously referred to as Padang Large morphotype) Rana cf. chalconota Inger & Iskandar, 2005: 138; Stuart et al., 2006: 473. Holotype FMNH 268580 (field no. 15864), an adult female from Limau Manis, 373 m (0°54′S / 100°28′E), Padang, West Sumatra, Indonesia. Collected in a disturbed forest 7.vii.2001, by Djong Hon-Tjong and David Gusman. Paratypes FMNH 268572, 268578-79, two adult females and one juvenile collected at same site and elevation as holotype on 3.vii. and 7.vii.2001; FMNH 268573-77, 268581-83, four adult males, four adult females from same locality as holotype, but at 405 m on 4.vii. and 10–11.vii.2001; FMNH 268584, 268587-88 two adult males, one adult female from Padang Jernih (0°52′S / 100°28′E) 255–340 m, Padang, West Sumatra, 26.vii.2001; FMNH 268585-86, one adult male, one juvenile from Sikayan Ubi (0°53′S / 100°27′E) 292 m, Padang, West Sumatra, 23.vii.2001. All with same collectors as holotype. Etymology Specific name from rufus, L., meaning reddish, and pes, L., meaning foot, referring to the reddish tinge on the underside of the webbing in life. Diagnosis A large form of the Rana chalconota group, adult females 46–64 mm SVL, males with nuptial pads 44–48 mm. Dark spots present on back. Nuptial pad of males not constricted. Humeral gland of males visible only by dissection. Tympanum relatively small, TYM/ SVL of females usually Description Habitus moderately slender, head slightly wider than trunk, legs long. Head triangular, slightly longer than broad; snout obtusely pointed, projecting slightly beyond lower jaw, longer than diameter of eye; nostril on side of snout, closer to tip of snout than to eye; canthus angular, not constricted; lores vertical, concave; interorbital wider than upper eyelid and wider than internarial; pineal body faintly visible between anterior corners of upper eyelids; tympanum distinct, about diameter of eye in females, slightly larger in males, inner portion slightly depressed; vomerine teeth in short, oblique groups, distance between groups equal to distance from choanae. Fingers long, length of third finger equal to distance from rear of eye to nostril; without webbing; second and third fingers with narrow, movable fold of skin on medial margins; tips of three outer fingers with wide discs, that of third finger three-quarters or more the diameter of the tympanum in females, disc of first finger about half width of disc of second finger, all discs with circummarginal grooves; subarticular tubercles conspicuous; bases of third and fourth fingers with one or two supernumerary tubercles, base of second finger with one; finger lengths 3> 4> 2> 1. Tips of toes expanded into discs smaller than those of fingers, but with circummarginal grooves; webbing extensive, to base of discs on lateral margins of first three toes and on medial margin of fifth, medial edge of fourth toe fully webbed to just beyond the distal subarticular tubercle; narrow dermal ridge along medial edge of distal joint of first toe and along lateral edge of distal joint of fifth toe; a low, oval inner metatarsal tubercle, shorter than distance to subarticular tubercle of first toe; a distinct, round outer metatarsal tubercle. Skin of back granular in females, in males granules tipped with colourless asperities or spinules; similar spinules present on lores in some males, the variation probably an artefact of preservation; a distinct, low dorsolateral fold; rear of abdomen rugose, rest of venter smooth. Males with paired vocal sac openings on floor of mouth. Whitish, velvety nuptial pad on dorsal and medial surfaces of first finger, not constricted. The humeral gland is detectable only by cutting and folding back the skin of the upper arm. Colour in preservative medium brown dorsally and on sides; side of head dark brown, upper lip white; dorsal surfaces with small dark spots; ventral surfaces of body whitish, unmarked; dark crossbars visible on hind limb only in a few individuals; ventral surface of webbing reddish, the colour fading in preservative. Measurements (mm) of holotype: SVL 62.0, tibia 34.3, head width 18.9, head length 23.2, tympanum diameter 4.8, width of disc of third finger 4.1. Variation Females 53.8–64.4 mm, mean 60.58 ± 1.55 mm (N = 6); males 43.7–48.4 mm, mean 45.36 ± 0.51 mm (N = 8). In the following data on body proportions, N = 7 for both sexes. T / SVL 0.537 –0.591, median 0.560 (N = 12), HW/ SVL of females 0.267 –0.312, of males 0.287 –0.309, HL/ SVL of females 0.360 –0.397, of males 0.370 –0.389, TYM/ SVL of females 0.065 –0.077, of males 0.097 –0.108; DF3/ SVL 0.053 –0.072, median 0.062 (N = 11). Comparisons Rana rufipes differs conspicuously from the form with which it co-occurs in West Sumatra, R. parvaccola (see below), in size, coloration of the webbing (Inger & Iskandar, 2005), relative size of the tympanum (TYM / SVL) and width of the disc of the third finger (DF 3/ SVL) (see Tables 2 and 6). The uncorrected pairwise sequence divergence between R. rufipes and the cooccurring R. parvaccola (see below) is 14.75–14.93% (Table 4). Rana rufipes is one of the largest members of this species group, with males larger than those of any other form and females larger than those of any other except the Javan species (Tables 2 and 6). This new species has the relatively smallest tympanum in the group, differing from all except males of R. labialis in TYM/SVL (Tables 2 and 6). It is also the only member of the group in which the ventral surface of the webbing is reddish., Published as part of Inger, Robert F., Stuart, Bryan L. & Iskandar, Djoko T., 2009, Systematics of a widespread Southeast Asian frog, Rana chalconota (Amphibia: Anura: Ranidae), pp. 123-147 in Zoological Journal of the Linnean Society 155 (1) on pages 137-138, DOI: 10.1111/j.1096-3642.2008.00440.x, http://zenodo.org/record/5445984, {"references":["Inger RF, Iskandar DT. 2005. A collection of amphibians from West Sumatra, with description of a new species of Megophrys (Amphibia: Anura). The Raffles Bulletin of Zoology 53: 133 - 142.","Stuart BL, Inger RF, Voris HK. 2006. High level of cryptic species diversity revealed by sympatric lineages of Southeast Asian forest frogs. Biology Letters 2: 470 - 474."]}