Lachesis puniceus Craspedocephalus puniceus Kuhl, 1824 Boulenger (1896: 561 1920: 291 De Rooij (1917: 286) Trimeresurus puniceus : Lidth de Jeude (1922: 252) Mertens (1934: 696) Kopstein (1938: 133) De Haas (1950: 610) Hoge & Romano (1974: 151) David & Vogel (1996: 165 David & Ineich (1999: 289) Orlov et al. (2002a: 194) Leong et al. (2003: 168 Trimeresurus cf. puniceus Trimeresurus cf. puniceus Trimeresurus cf. puniceus Trimeresurus puniceus A revision of the Trimeresurus puniceus-complex (Serpentes: Viperidae: Crotalinae) based on morphological and molecular data David, Patrick Vogel, Gernot Vijayakumar, S. P. Vidal, Nicolas Zootaxa 2006 2006-08-14 1293 1 1 78 [264,581,483,510] Reptilia Viperidae Trimeresurus Animalia Squamata 32 33 Chordata species puniceus  ( Figs. 12–14)     Lachesis puniceus(non  Craspedocephalus puniceus Kuhl, 1824):  Boulenger (1896: 561,  1920: 291);  De Rooij (1917: 286).     Trimeresurus puniceus: Lidth de Jeude (1922: 252);  Mertens (1934: 696)?;  Kopstein (1938: 133)?;  De Haas (1950: 610);  Hoge & Romano (1974: 151);  David & Vogel (1996: 165, Fig. 7);  David & Ineich (1999: 289);  Orlov et al.(2002a: 194);  Leong et al.(2003: 168, 171).   FIGURE 12.  Trimeresuruscf. puniceus. Specimen from Anamba Island (USNM 26544). General view. Photograph by Gernot Vogel. This chresonymy includes only citations based on the specimen from Mt. Kerinci, definitely identified as  Trimeresuruscf. puniceusas conceived here, on other specimens from Sumatera Barat Province(Alahanpanjang [ Kopstein, 1938] and Singkarak [ Mertens, 1934]), and specimens from Natuna Islands and Anamba Islands, which are all tentatively and somewhat arbitrarily referred to as  Trimeresuruscf. puniceus. General references merely citing  Trimeresurus puniceusfrom Sumatrawithout precision are not included.   Material ( 5 specimens).   INDONESIA. SumatraIsland. BMNH 1915.12.2.43 (female), “Sungei Kumbang, Sumatra,  4700 ft.”, now Sungai[Stream] Kumbang, a stream on slopes of Mt. Kerinci, Provinceof Sumatera Barat. Mentawai Archipelago. USNM 31696(female), Pulau Pagai Utara( North Pagai Island), Provinceof Sumatera Barat, Mentawai Archipelago. Natuna Islands. BMNH 95.5.1.100 (female), “Pulu Laut, Natuna Islands”, now Pulau Laut, Kepulauan Natuna; BMNH 95.5.1.101 (male), “Great Natuna Island”, now Pulau Natuna Besar, Kepulauan Natunas. Natuna Islands. USNM 26544(male), Pulau Siantan, Kepulauan Anambas( Anambas Islands).   FIGURE 13.  Trimeresuruscf. puniceus. Specimen from Anamba Island (USNM 26544). Lateral view of head, left side. Photograph by Gernot Vogel.  Taxonomic comments. These five specimensappear more or less morphologically close each from the other. Inmultivariate analyses, they form Cluster II, at the exception of the sole specimen from the Mentawai Archipelago( OTU 9) which is the most differentiated. Incontrast, multivariate analyses do not clearly separate specimens of OTU 3 ( Natuna+ Anamba Islands) from the sole available specimen of OTU 2 from West Sumatra, although there are morphological differences (see below). Our limited available material does not allow us to ascertain the relationships of these specimens, nor their relationships with specimens from OTU 2 (Southern Sumatra). Furthermore, if the molecular analyses clearly show that Sumatran specimens show a high genetic divergence (distance between them of 8.5 %), these results were obtained from two specimenswith no precise locality. Atthe present time, we cannot determine if we are in presence of two different taxa in southern and western Sumatra.  Leong et al.(2003)and Grismer et al.(2006)pointed out that Natuna and Anamba Archipelago, along with the islands of the Seribuat Archipelago, have phylogenetic relationships with some islands of the Sunda Shelf. Pending the collect of further specimens from western Sumatraand other islands, we refrain to conclude on the taxonomic position of these animals, and we just refer it to as  Trimeresuruscf. puniceus. We purposely adopt a conservative approach in not trying to define weakly supported new taxa on such a limited sample, being fully aware that these five specimensmay well represent two or even three distinct taxa. The taxonomy of the  T. puniceus­group will be addressed in a subsequent paper. This unnamed taxon appears in the key given below.   Diagnosis.These specimens are referred to as  Trimeresuruscf. puniceuson the basis of: (1) an overall brown coloration, with darker crossbands; (2) a distinctly projected and raised snout, strongly obliquely truncated when seen from the side; (3) internasals slightly extending out, weakly bilobate and sligthly raised; (4) 2 ndsupralabials not bordering the loreal pit, topped by a prefoveal scale; and (4) 3–4 small, distinct supraoculars. Characters (3) and (4) separate  Trimeresuruscf. puniceusfrom members of the  T. borneensis­group.   Description and variation. The longest examined specimen is a female, BMNH 1915.12.2.43, which is 609 mmlong (SVL 518 mm, TaL 91 mm). Morphology of head and body as described above for  Trimeresurus puniceus, with, in all specimens, a triangular head, wide at its base, thick and flattened. Snout distinctly projected and raised anteriorly, strongly obliquely trunctated when seen from the side, with a distinct canthus rostralis, subrectangular with slightly bilobate and upturned nasals. Most important characters are given in Table 9. Other main head scalation features include: two normal, upturned internasals on each side, separated by 1 scale in the three specimensof OTU 3, 2 scales in others; 2–3 small postoculars; 3 or 4 small, narrow supraoculars on each side, more or less pyramidal, extending out of the margin of the head and raised in OTU 2, strongly erect in OTU 9, and flat ( 2 specimens) or convex ( 1 specimen) in OTU 9; 6–8 slightly enlarged scales on upper snout surface, smooth, flat and juxtaposed; occipital and temporal scales smooth or weakly keeled; 9–11 SL; 3 rdSL separated from the subocular by 1 or 2 scales; 4 thSL and 5 thSL separated from subocular by 2 scales on each side in all specimens; 11–14 IL. In preservative, the background colour of these specimens is dark brown or dark greyish­brown, with irregular dark brown or nearly black dorsal markings, as in  T. puniceus. Male from Anamba Islands (USNM 26544) is dark seagrey with dark grey dorsal blotches. Postocular streak present, although quite subdued. Venter dark brown or dark grey, depending on specimens.  Comparison with other species.As a whole,  Trimeresuruscf. puniceusdiffers essentially from  T. puniceusby (1) a low number of ventral scales in females, 141–144 vs. 153–163 infemales of  T. puniceus, and (2) supraocular scales more raised and extending out from the lateral side of the head. All specimens of Cluster II are morphologically differentiated from other specimens of the  T. puniceus­group from Javaand the provinces of Bengkuluand Lampangof South Sumatra(Cluster I), referred above to as  Trimeresurus puniceus(see below).  The two specimensfrom West Sumatraand the adjacent North Pagai Islandespecially show a lower number of ventral scales. Thosefrom the Natunaand Anamba Islandsshow also a lower number of ventrals and are differenciated by flat supraoculars.  Range ( Fig. 4). INDONESIA.  SumatraIsland.  Sumatera Barat Province: Alahanpanjang ( Kopstein, 1938), Mt. Kerinci ( Boulenger, 1920; examined specimen), Singkarak ( Mertens, 1934). Mentawai Archipelago. Pulau Pagai Utara (North Pagai Island) (examined specimen). Kepulauan Natuna (Natuna Islands). Pulau Natuna Besar (Bunguran, or Great Natuna Island) and Pulau Laut (Laut Island) (examined specimens). Kepulauan Anambas (Anamba Islands). Pulau Siantan (Siantan Island; examined specimen).   TABLE 9.Main characters of specimens referred to as  Trimeresuruscf. puniceus.    POPULATION TaL/TL VEN SC MSR Cep SupOc shape  ♂ ♀ ♂ ♀ ♂ ♀  OTU 2 (W. Sumatra) BMNH 1915.12.2.43 0.149 141 43 21 12 raised  OTU 3 (Natuna & Anamba Is.)  BMNH 95.5.1.101 USNM 26544 0.170 0.168 155 163 53 55 21 19 13 12 flat flat  BMNH 95.5.1.100 0.131 148 ..... 21 11 convex  OTU 9 (Pagai Is.) USNM 31696 0.136 144 40 21 14 erect    Trimeresurus puniceus(n = 45) 0.150– 0.171 0.128– 0.151 155–167 153–163 44–59 41–49 19–23 12–15 raised In the southern part of its range,  Trimeresuruscf. puniceusmay occur in sympatry with  Trimeresurus puniceus, whereas, in the north, it may occur in sympatry with  Trimeresurus andalasensis(see below), also known from the Padang highlands. Specimens recorded by Matsui et al.(1984)from Mt. Gadut, Sumatera Barat Province, are here referred to as  Trimeresurus andalasensis  spec. nov.(see below) and not to  T. puniceusor  T.cf. puniceusas suggested in David & Vogel (1996; as  T. puniceus) on the basis of ventral scale counts. We cannot identify to which species should be referred the specimen from Padang, Province of Sumatera Baratrecorded in Hubrecht (1887; as Trigonocephalus puniceus).   Biology.According to Boulenger (1920), specimen BMNH 1915.12.2.43 was collected on slopes of Mt. Kerinci at 4700 feet. We have no data for other specimens.   Comments.None of the specimens referred here to  Trimeresuruscf. puniceushas been depicted in the literature. Specimens from Indrapura, Sumatera Barat Province, are depicted in Gumprecht & Tepedelen (1999: 26: Figs. 1–3; 28: Fig. 7; 29: Figs. 8–9); they are identical in pattern and coloration with specimens from Bengkulu. As these authors did not provide detailed scale counts, we cannot identify these specimens. Nevertheless, they might be conspecific with the specimen from Mt. Kerinci identified here as  Trimeresuruscf. puniceus. BMNH, USNM Indonesia Sungai 1433 Mt. Kerinci Kumbang 33 34 BMNH 1915.12, USNM 31696, BMNH 95.5, USNM 26544 1 Sumatera Utara OTU In & Cluster II & Mentawai Archipelago In 34 35 Anamba Islands Natuna 33 34 OTU 9 1 Sumatera Barat At Sumatra 34 35 1 Sumatera Utara North Pagai Island Natuna Those 35 36 1 Sumatera Barat