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A new species of Calamaria (Squamata: Colubridae) similar to C. ceramensis de Rooij, 1913, from the Banggai Islands, east of Sulawesi, Indonesia ANDRÉ KOCH1,5, EVY ARIDA2, JIMMY A. MCGUIRE3, DJOKO T. ISKANDAR4 & WOLFGANG BÖHME1 1
Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, 53113 Bonn, Germany Museum Zoologicum Bogoriense, Jl. Raya Bogor km 46, 16911 Cibinong, Indonesia; current address: Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, 53113 Bonn, Germany 3 Museum of Vertebrate Zoology and Department of Integrative Biology, 3101 Valley Life Sciences Building, University of California, Berkeley, CA 94720-3160, USA 4 School of Life Sciences and Technology, Institut Teknologi Bandung, 10 Jalan Ganesa, Bandung 40132, Indonesia 5 Corresponding author. E-mail:
[email protected] 2
Abstract A new species of reed snake, genus Calamaria, is described from the Banggai Islands off the east coast of Central Sulawesi, Indonesia. The new species, which is phenotypically similar to C. ceramensis from the central Moluccas, is characterised by the absence of preocular scales, five supralabials (third and fourth entering orbit), five infralabials (the first meet behind the mental), mental scale separated from the anterior chin shields, five scales surrounding the paraparietal, (most probably) seven modified maxillary teeth, a high number of ventral scales, a short, thick tail which is slightly tapering to a point, a single anal plate, as well as by a characteristic colour pattern of a pale brown dorsum dotted with darker spots, a pale collar behind the head, and a pale and dark lateral zigzag pattern along the body. Calamaria banggaiensis sp. nov. represents the fifty-eighth known species of Calamaria and the twelfth species of this genus recorded from the Sulawesi region. The new taxon is the only species of reed snake recorded from the Banggai Islands. It is known only from two adult specimens, a male and a female. For phenotypic comparison with the new species, both syntypes of C. ceramensis de Rooij, 1913, are depicted herein for the first time. An updated key to the Calamaria species of Sulawesi and its satellite islands is presented. Key words: Calamaria banggaiensis sp. nov., Indonesia, Moluccas, Seram, Sulawesi region, Wallacea
Abstrak (Bahasa Indonesia) Telah dideskripsi satu species baru ular pada marga Calamaria yang berasal dari Kepulauan Banggai yang termasuk di dalam wilayah provinsi Sulawesi Tengah, Indonesia. Species baru yang secara fenetis mirip dengan C. ceramensis (dari Kepulauan Maluku Tengah) ini, dideskripsikan dengan ciri-ciri sebagai berikut: tidak terdapat sisik preokular, terdapat lima sisik supralabial (sisik ketiga dan keempat berhadapan langsung dengan mata), lima sisik infralabial (sisik pertama bertemu di belakang sisik mental), sisik mental terpisah dari sisik dagu (chin shields) anterior, dan lima sisik mengitari sisik parietal; (kemungkinan besar) 7 gigi yang termodifikasi pada rahang atas, jumlah sisik ventral yang banyak (198 buah), terdapat 20 sisik subkaudal yang berpasangan, ukuran tubuh kecil (panjang total 201 mm), ekor pendek dan tebal meruncing, satu buah sisik anal, serta pola warna yang khas dengan dasar coklat terang dan totol-totol gelap, sisik nukal berwarna terang di bagian belakang kepala, dan motif zigzag gelap-terang sepanjang tubuh. Jenis baru ini adalah yang ke-58 dari semua jenis ular dalam marga Calamaria dan jenis ke-12 yang ditemukan di daerah Sulawesi dan sekitarnya. Jenis baru ini merupakan satu-satunya dalam marga Calamaria dari Kepulauan Banggai, berdasarkan pada dua spesimen tipe dewasa. Sebagai pembanding karakter fenotip untuk jenis baru ini, kedua sintipe dari C. ceramensis de Rooij, 1913, ditampilkan pada artikel ini. Selain itu, kunci determinasi untuk marga Calamaria dari Sulawesi dan pulau-pulau kecil di sekitarnya juga telah diperbaharui dalam artikel ini. Accepted by D. Gower: 3 Jun. 2009; published: 13 Aug. 2009
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Introduction The Asian colubrine genus Calamaria Boie, 1827, currently comprises fifty-seven recognised species. This group of fossorial snakes is distributed from China and the Ryu Kyu Islands (Japan) southwards through Vietnam, Cambodia, Laos, Myanmar, Thailand, Malaysia, the Indonesian Archipelago including the Greater Sunda Islands, Bali and Sulawesi, the Philippines, and as far as the Cocos (Keeling) Islands (Australia) (Inger & Marx 1965). The easternmost representative of the genus is C. ceramensis from the islands of Seram, Saparua and Ambon in the central Moluccas (Inger & Marx 1965). Due to the superficially homogeneous morphology of Calamaria species (all species have 13 dorsal scale rows and relatively few head scales) only four generic names have been proposed for these small to medium-sized snakes since the genus Calamaria was posthumously established by Heinrich Boie (1794–1827) in the year of his death (H. Boie in F. Boie 1827). In their monumental study of the genus Calamaria, Inger & Marx (1965) provided a comprehensive revision of the genus which, at that time, included 50 recognized species. Since that work, only six new taxa have been added. Four of these taxa are from mainland Southeast Asia: C. ingeri Grismer et al. (2004) from Pulau (= Island) Tioman, Malaysia, and C. lovii ingermarxorum Darevsky & Orlov (1992; new name combination according to Michels & Bauer 2004), C. thanhi Ziegler & Le (2005), and C. gialaiensis Ziegler et al. (2008) from Vietnam. Gillespie et al. (2005) reported two new Calamaria species from Pulau Buton, the largest of Sulawesi’s satellite islands. These were recently described as C. butonensis and C. longirostris by Howard & Gillespie (2007). In addition, two names, C. pfefferi Stejneger, 1901, and C. yunnanensis Chernov, 1962, from the Ryu Kyu Islands and China, respectively, were resurrected from synonymy (Ota 1982; Zhao & Adler 1993). Each of these ‘new’ Calamaria taxa has been discovered either from the margin of the geographic range of the genus, or from small islands. These findings suggest that the diversity of Asian reed snakes is better understood than is the case for most other amphibian and reptile genera in Southeast Asia, and suggests that newly discovered taxa are most likely to be encountered on remote islands and other poorly explored regions. One such remote and poorly explored locality is Kepulauan (= archipelago) Banggai, an island group located between the east coast of Central Sulawesi and the Sula Islands in the western Moluccas (Fig. 1). During field work on the islands of Banggai and Peleng we collected two specimens of Calamaria. Although Koch et al. (2007a) preliminarily allocated the Banggai specimen to C. cf. ceramensis, we have since made detailed morphological comparisons with the type specimens of C. ceramensis, leading us to the conclusion that these specimens represent a new species, which we describe herein.
Material and methods Herpetofaunal surveys on the islands of Banggai and Peleng were conducted between 22 and 30 August 2005 and between 15 and 18 September 2007, respectively. AK and JAM field numbers refer to the collections made by AK/EA and JAM/DTI during field work, respectively. Terminology and methodology for the description of the new taxon follow Inger & Marx (1965) and Grismer et al. (2004). All measurements from preserved specimens were taken by AK and JAM using a dial calliper to the nearest 0.1 mm except for snout-vent and tail lengths, which were recorded with a measuring tape to the nearest 1 mm. Relevant voucher specimens and type material of Sulawesian Calamaria species were examined for taxonomic comparisons (Appendix). Museum acronyms employed here follow Leviton et al. (1980). The holotype of the new species is stored in the herpetological collection of the Museum Zoologicum Bogoriense (MZB), Indonesia. Also, the paratype currently held at the Institut Teknologi Bandung (ITB) will ultimately be deposited in the MZB collection.
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FIGURE 1. Map of the Sulawesi region and the Moluccas. Banggai Island, the type locality of Calamaria banggaiensis sp. nov., located east off Peleng is indicated by an asterisk. Question marks denote islands from which Calamaria have not been reported, between the disjunct distribution ranges of C. banggaiensis sp. nov. and C. ceramensis in the Moluccas.
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 JAM8759) 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.
A NEW CALAMARIA FROM THE BANGGAI ISLANDS
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FIGURE 2. Holotype of Calamaria banggaiensis sp. nov. in life showing the characteristic dorsal pattern. Photo by André Koch.
FIGURE 3. Dorsal view of syntypes (ZMA 10083) of C. ceramensis de Rooij, 1913, from Seram Island, Moluccas. Photo by André Koch.
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FIGURE 4. Ventral view of the holotype of Calamaria banggaiensis sp. nov. (MZB Oph. 3230). Note the absence of a dark median stripe on the ventral surface of the tail. Photo by André Koch.
FIGURE 5. Ventral view of syntype ZMA10083a of Calamaria ceramensis illustrating the dark median stripe on the ventral surface of the tail. Photo by André Koch.
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. A NEW CALAMARIA FROM THE BANGGAI ISLANDS
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FIGURE 6. Scalation details of the head of the holotype of Calamaria banggaiensis sp. nov., dorsolateral view. Drawing by André Koch.
FIGURE 7. Lateral view of the head of the syntype (ZMA 10083b) of Calamaria ceramensis. Photo by André Koch.
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FIGURE 8. Dorsal view of the head of the holotype of Calamaria banggaiensis sp. nov. Photo by Evy Arida.
FIGURE 9. Dorsal view of the head of the syntype (ZMA 10083a) of Calamaria ceramensis. Photo by André Koch.
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
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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
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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. 2007a).
Discussion Calamaria banggaiensis is the fifty-eighth currently recognised species of Calamaria and the twelfth member of this genus recorded from the Sulawesi region sensu Vane-Wright (1991), though the only species of reed snake recorded from the Banggai Islands. In several characters (i.e. modified maxillary teeth, mental not A NEW CALAMARIA FROM THE BANGGAI ISLANDS
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touching anterior chin shields, absence of preocular scales, prefrontal touching first three supralabials, five scales surrounding the paraparietal and the presence of a light nuchal collar), C. banggaiensis is most similar (and perhaps most closely related) to C. ceramensis, the southeastern-most representative of calamarine reed snakes and the only other species previously known to occur east of Sulawesi. The discovery of C. banggaiensis on the Banggai Islands partially closes the wide gap between the disjunct distributions of C. ceramensis in the central Moluccas and its nearest congeners on Sulawesi by approximately 120 kilometres. Although Calamaria ceramensis was only rarely recorded in the past (e.g. by Smith & Procter, 1921), such that less than a dozen voucher specimens were available to Inger & Marx (1965), this snake species seems not to be rare on Seram. According to Edgar & Lilley (1987), C. ceramensis was encountered at any time of day or night and in various environments. The species was found in coastal and lowland forests as well as secondary and primary rain forest at 700 m altitude or above, and in human modified habitation such as plantations. Nevertheless, abundances of distinct Calamaria taxa may vary remarkably, as reported in a longterm field survey by Howard & Gillespie (2007) for two sympatric species (C. butonensis and C. longirostris, respectively) on Buton Island, off the southeastern peninsula of Sulawesi. This observation is perhaps explained by differing degrees of fossoriality exhibited by various species, with more fossorial species likely to be encountered less frequently during standard herpetological surveys. With respect to the biogeography of reed snakes, Inger & Marx (1965) hypothesized that the Sulawesi Calamaria fauna was established via two or three independent dispersal events from adjacent Borneo. They further suggested a single dispersal event to the east of Sulawesi, resulting in the highly disjunct C. ceramensis in the Moluccas. We can now propose that at least two over-water dispersal events were necessary to explain the distribution of Calamaria, because the Banggai Archipelago is separated from both Sulawesi and the Moluccan islands by deep ocean trenches. However, in the absence of a phylogenetic estimate for the genus, the direction of dispersal and source populations remain unclear. The occurrence of a Calamaria species on the Banggai Islands agrees with How & Kitchener’s (1997) hypothesis of a major zoogeographic division between the snake faunas of the Moluccas in the east and those of Sulawesi and the Lesser Sunda Islands in the west, as well as with a faunistic link at species level between the Banggai and Sula Archipelagos to the Lesser Sundas. Eleven out of twelve Sulawesian Calamaria species (except for C. virgulata) are endemics and the new described species may prove to be restricted to the Banggai Archipelago and possibly the Sula Islands further east (Fig. 1). Nevertheless, it remains unclear which species inhabit Pulau Buru, located between the Banggai Islands and Seram along the natural route of dispersal, as well as on other Moluccan islands that remain poorly explored. In recent years, systematic investigations of the long-neglected herpetofauna of Sulawesi have shown that the herpetological diversity of the region is far from completely known (Iskandar & Tjan 1996; de Lang & Vogel 2005; Gillespie et al. 2005; Koch et al. 2007a, b; McGuire et al. 2007; Linkem et al. 2008). Therefore, future fieldwork on the Banggai and Sula Islands will probably reveal more undescribed species, further detailing the evolutionary history and diversity of Calamaria and other amphibians and reptiles in the region.
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 2 3 4 -
Preocular absent ......................................................................................................................................................... 9 Preocular present ......................................................................................................................................................... 2 Mental not touching anterior chin shields .................................................................................................................... 3 Mental touching anterior chin shields........................................................................................................................... 5 Paraparietal surrounded by five scales and shields................................................................................ C. brongersmai Paraparietal surrounded by six scales and shields ........................................................................................................ 4 Eye diameter almost 1.5 times eye-mouth distance................................................................................. C. boesemani Eye diameter slightly greater than eye-mouth distance .............................................................................. C. virgulata
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First gular touching anterior chin shields ................................................................................................ C. acutirostris First gular not touching anterior chin shields .............................................................................................................. 6 Paraparietal surrounded by five scales and shields....................................................................................... C. muelleri Paraparietal surrounded by six scales and shields ....................................................................................................... 7 Tail thick, tapering abruptly at end .............................................................................................................C. virgulata Tail tapering gradually from base ............................................................................................................................... 8 Ventrals yellow, immaculate except for dark lateral tips ............................................................................. C. nuchalis Ventrals dark, yellowish on posterior edges only .............................................................................................. C. curta Mental touching anterior chin shields................................................................................................ C. apraeocularis Mental not touching anterior chin shields.................................................................................................................. 10 Four supralabials..................................................................................................................................... C. longirostris Five supralabials ......................................................................................................................................................... 11 Pale collar absent ...................................................................................................................................... C. butonensis Pale collar present....................................................................................................................................................... 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
Acknowledgements We are greatly indebted to Prof. Dr. Dedy Darnaedi (Research Center for Biology, Bogor, Indonesia) and Dr. Mulyadi (MZB, Indonesia) for their letters of support and to the Indonesian Institute of Sciences (LIPI) for research (AK: 2767/SU.3/KS/2005; JAM: 4224/SU.3/KS/2007) permits in support of our biodiversity sampling on Sulawesi and adjacent islands. AK and EA would like to express their gratitude to Pak Antok and to Ibu Yani and her husband Jérome Doucet (all Luwuk, Sulawesi) for their sincere support during field work in Central Sulawesi. Further, we thank Ibu Mumpuni (MZB), Franz Tiedemann and Heinz Grillitsch (all NMW, Austria), Raffael Winkler, and Ambros Hänggi and Urs Wüest (all NMB, Switzerland) for the permission to examine material under their care, and Ronald Vonk (ZMA, the Netherlands) for the loan of the syntypes of C. ceramensis. An earlier version of the present manuscript benefited from helpful comments by Thomas Ziegler (Cologne Zoo, Germany), Robert Neal (Brisbane, Australia), and Patrick David (Paris, France). “M & S Reptilien” (VS Weigheim, Germany) and Peter Hoch (Waldkirch, Germany) kindly provided equipment for field work of AK and EA on Sulawesi. Financial support was provided by a PhD grant to AK from the Evangelisches Studienwerk e.V. Villigst. Fieldwork of JAM and DTI was supported by the National Science Foundation (DEB 0328700 DEB and 0640967). This is publication No. 8 of the cooperation project on the herpetofauna of Sulawesi between the herpetological sections of the Zoologisches Forschungsmuseum A. Koenig (ZFMK), Germany, and the Museum Zoologicum Bogoriense (MZB), Indonesia.
References Boie, F. (1827) Bemerkungen über Merrem’s Versuch eines Systems der Amphibien. 1820. Marburg. Erste Lieferung: Ophidier. Isis von Oken, 20(10), 508–566. Darevsky, I.S. & Orlov, N.L. (1992) A new subspecies of the dwarf snake Calamaria lowi ingermarxi ssp. nov. (Serpentes, Colubridae) from southern Vietnam. Asiatic Herpetological Research, 4, 13–17. 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. de Rooij, N. (1913) Praeda itineris a L. F. de Beaufort in Archipelago indico facti annis 1909–1910. III. Reptilien. Bijdragen tot de Dierkunde, 19, 15–29. Edgar P.W. & Lilley, R.P.H. (1987) Herpetofauna survey of Manusela National Park. In: Edwards I.D., MacDonald A.A. & J. Proctor (Eds.), Natural History of Seram. Andover, Intercept, pp. 131–142. Gillespie, G., Howard, S., Lockie, D., Scroggie, M. & Boeadi (2005) Herpetofaunal Richness and Community Structure of Offshore Islands of Sulawesi, Indonesia. Biotropica, 37(2), 279–290.
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Appendix. Specimens examined Calamaria acutirostris (2)—NMB 1686 (lectotype) from Loka, near Bonthain (= Bantaeng), Southwest Sulawesi; NMW 16714 from Loka, Southwest Sulawesi. C. brongersmai (1)—NMB 1707 (paratype) from Lake Posso, Danau Poso, Central Sulawesi. C. butonensis (2)—MZB Oph. 2976 (paratype) from Labundo Bundo, Buton Island, South Sulawesi; MZB Serp. 2985 (paratype) from Labundo bundo, Buton Island, South Sulawesi. C. ceramensis (2)—ZMA 10083[2] (syntypes) from Honitetu, West Seram. C. longirostris: MZB Oph. 2849 (paratype) from Labundo Bundo, Buton Island, South Sulawesi. C. muelleri (5)—ZSM 545/1920 from Bonthain (= Bantaeng), Southwest Sulawesi; NMW 16691 from Bonthain (= Bantaeng), Southwest Sulawesi; MZB Oph. 2344 from Gowa, South Sulawesi; MZB Oph. 2345 from Gowa, South Sulawesi; MZB Oph. 2346 from Gowa, South Sulawesi. C. nuchalis (2)—NMW 16715 from Bua Praeng (in error for Mt. Bawakaraeng), Southwest Sulawesi; MZB Oph. 1780, Tomado, Lake Lindu, Central Sulawesi. C. virgulata (3)—NMB 1695 (holotype of Calamaria collaris) from Tomohon, North Sulawesi; NMW 16716-1 and NMW 16716-2 from Bua Praeng (in error for Mt. Bawakaraeng), Southwest Sulawesi; C. cf. virgulata (1)—ZMH R-08624 from Rurukan, North Sulawesi. Calamaria sp. (1) —MZB Oph. 3240 (AK0132) from Biak, near Luwuk, Central Sulawesi.
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