Calamaria banggaiensis, Koch, André, Arida, Evy, Mcguire, Jimmy A., Iskandar, Djoko T. & Böhme, Wolfgang, 2009

Koch, André, Arida, Evy, Mcguire, Jimmy A., Iskandar, Djoko T. & Böhme, Wolfgang, 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, pp. 19-30 : 21-27

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.

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. 2007a).

Figures

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.

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.

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 ZMA 10083 a of Calamaria ceramensis illustrating the dark median stripe on the ventral surface of the tail. Photo by André Koch.

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 10083 b) of Calamaria ceramensis. Photo by André Koch.