A revision of the Trimeresurus puniceus-complex (Serpentes: Viperidae: Crotalinae) based on morphological and molecular data Author David, Patrick Author Vogel, Gernot Author Vijayakumar, S. P. Author Vidal, Nicolas text Zootaxa 2006 2006-08-14 1293 1 1 78 https://biotaxa.org/Zootaxa/article/view/zootaxa.1293.1.1 journal article 10.11646/zootaxa.1293.1.1 1175­5334 5070879 Trimeresurus cf. 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. Trimeresurus cf. 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 Trimeresurus cf. puniceus as 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 Trimeresurus cf. puniceus . General references merely citing Trimeresurus puniceus from Sumatra without precision are not included. Material ( 5 specimens ). INDONESIA . Sumatra Island . BMNH 1915.12 .2.43 (female), “Sungei Kumbang, Sumatra , 4700 ft. ”, now Sungai [Stream] Kumbang , a stream on slopes of Mt. Kerinci , Province of Sumatera Barat . Mentawai Archipelago . USNM 31696 (female), Pulau Pagai Utara ( North Pagai Island ), Province of 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. Trimeresurus cf. puniceus . Specimen from Anamba Island (USNM 26544). Lateral view of head, left side. Photograph by Gernot Vogel. Taxonomic comments. These five specimens appear more or less morphologically close each from the other. In multivariate analyses, they form Cluster II , at the exception of the sole specimen from the Mentawai Archipelago ( OTU 9 ) which is the most differentiated. In contrast, 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 specimens with no precise locality. At the 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 Sumatra and other islands, we refrain to conclude on the taxonomic position of these animals, and we just refer it to as Trimeresurus cf. 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 specimens may well represent two or even three distinct taxa. The taxonomy of the T. puniceus ­group will be addressed in a subsequent paper. This unnamed taxon appears in the key given below. FIGURE 14. Trimeresurus cf. puniceus . Specimen from Anamba Island (USNM 26544). Dorsal view of head. Photograph by Gernot Vogel. Diagnosis. These specimens are referred to as Trimeresurus cf. puniceus on 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 nd supralabials not bordering the loreal pit, topped by a prefoveal scale; and (4) 3–4 small, distinct supraoculars. Characters (3) and (4) separate Trimeresurus cf. puniceus from members of the T. borneensis ­group. Description and variation . The longest examined specimen is a female, BMNH 1915.12.2.43, which is 609 mm long (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 specimens of 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 rd SL separated from the subocular by 1 or 2 scales; 4 th SL and 5 th SL 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, Trimeresurus cf. puniceus differs essentially from T. puniceus by (1) a low number of ventral scales in females, 141–144 vs. 153–163 in females 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. puniceus ­group from Java and the provinces of Bengkulu and Lampang of South Sumatra (Cluster I), referred above to as Trimeresurus puniceus (see below). The two specimens from West Sumatra and the adjacent North Pagai Island especially show a lower number of ventral scales. Those from the Natuna and Anamba Islands show also a lower number of ventrals and are differenciated by flat supraoculars . Range ( Fig. 4 ). INDONESIA . Sumatra Island . 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 Trimeresurus cf. 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, Trimeresurus cf. puniceus may 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. puniceus or T. cf. puniceus as 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 Barat recorded 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 Trimeresurus cf. puniceus has 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 Trimeresurus cf. puniceus .