1 Bonn zoological Bulletin Volume 60 Issue 1 pp. 17-24 Bonn, May 201 Description of a new species of the genus Dendrelaphis Boulenger, 1890 from Myanmar (Squamata: Serpentes: Colubridae) Gemot Vogel>*& Johan van Rooijen2-3 1 Society for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, Germany; E-mail: [email protected] 2 Netherlands Centre for Biodiversity Naruralis, section Zoological Museum Amsterdam, Mauritskade 61, AD 092 Amsterdam, The Netherlands 1 3 Tulpentuin 313, 2272 EH, Voorburg, The Netherlands; E-mail: [email protected] *corresponding author. Abstract. The population systematics ofthe colubrid snake so far referred to as Dendrelaphisgorei (Wall, 1910) were investigatedby carrying out a multivariate analysis ofgeographic variation. The results reveal the existence oftwo dis- tinct and apparently disjunct phenotypes. These phenotypes are assumed to represent independent evolutionary lineag- es. ThefirstlineagecorrespondswithD. goreiwhich inhabits theAssamValleyandadjoiningHimalayanfoothills. How- ever, the name Dendrophisgorei Wall, 1910 is actually ajunior synonym ofDendrelaphis biloreatus Wall, 1908. Con- sequently, the latter name is adopted for this lineage. The second lineage, described in this paper as a new species, in- habits the South Myanmar lowlands and neighbouring mountain chains. It is distinguished fromD. biloreatus on the ba- A sisofits higherventral count(203-212versus 190-199), itsundividedanal shield(dividedin biloreatus) anditshigh- eraverage number ofanteriortemporal shields (usually 2 versus usually 1). Key words. Dendrelaphis sp. n.,Dendrelaphisgorei syn. n., Dendrelaphis biloreatus, Myanmar, analysis ofgeograph- ic variation. INTRODUCTION The colubrid snakes ofthe genusDendrelaphis Boulenger, Forest Department, Ministry ofForestry, Myanmar (e.g. 1890 are widely distributed, ranging from Pakistan in the Wogan et al. 2008). In this paper, we investigate the pop- West to the northern and eastern coast ofAustralia in the ulation systematics ofD. goreibyperforming a multivari- Eastand South and to southernChina in theNorth (Ziegler ate analysis ofgeographic variation based on morpholog- & Vogel 1999). Members ofthe genus Dendrelaphis are ical data. Furthermore, the nameDendrelaphis biloreatus slender, diurnal species that are predominantly arboreal Wall, 1908 is evaluated. Smith (1943) synonymized D. and feed mainly on lizards and amphibians. biloreatus withD. gorei (Wall 1910) as he could not find any differences betweenthe two species apart fromthe di- & Boulenger(1894), Wall (1921), Meise Henning (1932), vided loreal shield in D. biloreatus. However, according Mertens (1934) and Smith (1943) have in turn worked on to the rule ofpriority (Art. 23, ICZN, 1999), the nameD. the systematics ofthis genus. Nevertheless, their cumu- biloreatus should be adopted for this species. lative effort did not result in an unambiguous and com- plete taxonomy ofthis genus, a fact that was underlined by the recent descriptions and revalidations of several MATERIALS AND METHODS species (Vogel & Van Rooijen 2007, 2008, 2011; Van Rooijen & Vogel 2008a, 2008b, 2008c, 2009). Eleven museum specimens were examined for this study. In addition, data on three additional specimens were tak- Dendrelaphisgorei (Wall, 1910) has been reported to oc- en from Wall (1908, 1910). Foreach examined specimen, cur inNortheast India and Myanmar. It is one ofthe least 23 characters including aspects of colour pattern, body known members ofits genus and is scarcely represented proportions and scalation were recorded (Table 1). Eye- in museum collections. In recent years however, new ma- diameter and distance eye-nostril were measured with a terial hasbecome availablethroughthe MyanmarHerpeto- slide calliperto the nearest 0.1 mm. These measurements logical Survey, a collaborative effort of the California were made on the left andright side andwere subsequent- Academy of Sciences, Smithsonian Institution and the ly averaged. Snout-vent length was measured to the pos- ) IN Gemot Vogel & Johan van Rooijen terior margin ofthe anal plate by marking the length on Geographic coordinates were taken directly from the field a piece ofstring and subsequently measuring the position notes or were obtained by translating locations to coordi- ofthe mark to the nearest 0.5 cm. Tail-length was meas- nates. Relevant variables were included in a Principal ured to the nearest 0.5 cm by straightening the tail against Components Analysis (PCA, e.g. Cramer 2003) in order a ruler. The number ofventrals was counted using Dowl- to reduce the dimensions of the dataset. The resulting ing's method (1951). Subcaudals were counted on one PCA-scores were then plotted against longitude and lati- side, the terminal scute was excluded. The first sublabial tude in order to visualize the pattern ofgeographic vari- was defined as the scale that starts between the posterior ation. Confirmatory geographic analyses ofthe PCA-score chin shield andthe infralabials and that borders the infral- were carried out by Analysis ofCovariance (ANCOVA; abials (see Peters 1964, fig. 7). The last infralabial was Maxwell & Delaney 1990; Norusis & SPSS 1993) using defined as the infralabial still covered completely by the longitude and latitude as covariates andphenotype as fac- last supralabial. The posterior most temporal scales were tor. Normality ofthe PC-scores was first testedby means defined as the scales ofwhich more than halfofthe area ofKolmogorov-Smirnov tests. lies in front ofan imaginary line that runs from the apex ofthe last supralabial to the posterolateral corner ofthe All statistical analyses were carried out with the software parietal. SPSS (2006; SPSS for Windows. Release 14.0.2. Chica- go: SPSS Inc.). Table 1. List ofmorphometric, scalation and coloration characters used in this study and their abbreviations. Abbreviation Character Morphometries EYED Horizontal diameter ofthe eye EYEN Distance from centre ofthe eye to posterior border ofthe nostril TAIL Tail-length WSNOUT Width ofthe snout measured at the position ofthe nostrils LHEAD Head-length measured from the tip ofthe snout to the rear ofthejaw SVL Snout-vent length Scalation VENT Number ofventrals SUBC Number ofsubcaudals DOR1 Number ofdorsal scale rows 1 head-length behind the head DOR2 Number ofdorsal scale rows at the position ofthe middle ventral DOR3 Number ofdorsal scale rows 1 head-length before the tail SUBL Number ofinfralabials touched by the first sublabial (L+R) SL1 Number ofsupralabials (L+R) SL2 Number ofsupralabials touching the eyes (L+R) LOR Number ofloreals (L+R) INFR Number ofinfralabials (L+R) ATEMP Number ofanterior temporals (L+R) TEMP Number oftemporals (L+R) POC Number ofpostoculars (L+R) VERT Vertebral scales smaller than (0) or larger than (1) scales ofthe first dorsal row AN Anal shield divided (0) or undivided (1) Coloration TSTRIPE Postocular stripe absent (0), rudimentary (1), present (2) LSTRIPE Ventrolateral stripe absent (0), present (1 BMNH The collected data were used to carry out an analysis of Material examined. S-037, Guwahati, Assam; BMNH geographic variation. The objective of this analysis was 1940.3.4.24, Assam; 1940.3.4.25, Samagooting, to enable differentiation between clinal variation andphe- Assam; BMNH 1946.1.10.30, Jaipur, Assam (syntype netic discontinuities, the latterbeing evidence for lineage Dendrophis gorei Wall, 1910); CAS 208429, Myanmar, separation (e.g. Lenk & Wiister 1999; De Queiroz 2007). Bago Division, BagoYoma, SeinYay Camp, 18 51 21.636 Bonn zoological Bulletin 60 (1): 17-24 ©ZFMK A new Dendrelaphis from Myanmar 19 N, 96 10 21.324 E; CAS 21 1939, Myanmar, Ayeyarwady RESULTS Mwe Division, vie HaiikVillage (16 16 39.2N, 9445 32.5 E); CAS 222114-222115, Myanmar: Bago Division,Aok Statistics TwinTownship, Ka Baung Reserve, SeinYe Camp, Block 120, 18 51 16.1 N, 96 10 23.6 E; CAS 222339, Myan- Visual inspection ofthe data suggested a striking differ- mar: Chin State, Min Dat Township, Min Dat Township, entiation between specimens fromAssam (NE India) and Nat Ma Taung National Park, 2121 14.9 N, 93 56 08.3 Northwest Myanmaron the one handand specimens from E; CAS 234880, Myanmar,: Chin State, MindatTownship, the remainder ofMyanmar on the other hand. One phe- MindatDistrict, 21 2643.5 N, 94 00 24.0 E; CAS 244037, notype was characterized by a high number of ventral Myanmar, Sagaing Division, Leha and Khante township counts, an undivided anal shield and usually two anteri- boundary, Nana Sa Laing camp, 26 07 23.1 N, 95 32 24.6 or temporal shields whereas the other was characterized by a lownumberofventrals, adivided anal shield andusu- ally one anterior temporal shield. These characters were included in a PCA. The first component explained 92% Museum abbreviations. BMNH: Natural HistoryMuse- ofthe total variance. This demonstrates that these char- um, London, Great Britain. CAS: Collection ofthe Cali- acters covary strongly and thus form a suite ofcharacters fornia Academy of Sciences, USA. S: Saibal Senguptas that sharply differentiates between the two phenotypes. Collection, Guwahati, Assam The individual scores on this component were plotted 1.0- 0.5- U o.o 0- 45' -1.0- 16 94 93 Situ de 92 28 Fig. 1. PCA-scores based on a PCA ofthe characters VENT, AN andATEMP against longitude and latitude. 1 Phenotype 1, 2 phenotype 2, 2b= type D. biloreatus Wall, 1908; 2a = syntype D. gorei Wall, 1910. Bonn zoological Bulletin 60 (1): 17-24 ©ZFMK 2(1 Gemot Vogel & Johan van Rooijen & against latitude and longitude to visualize the pattern of drelaphisproarchos (Wall, 1909) (Vogel van Rooijen geographic variation. In figure data-points correspon- 2011). We examined more than 600 museum specimens 1, ding with the type ofDendrelaphis biloreatus and one of ofthis genus, representing 24 species and subspecies. On- the two syntypes ofDendrophis gorei are indicated for a ly a single specimen (beside those identified as D.proar- later discussion ofthe appropriate nomenclature. A phe- chos and the material presented in this study) was found netic discontinuity is in evidence, the difference in PCA- to have an undivided anal shield. Second, the two pheno- scores between the phenotypes being highly significant types differ substantially in the number ofventral scales, (ANCOVA,df=l,ll; PO.0001). Subsequent inclusion in the difference being 19 on average (211 in phenotype 1, the model oflongitude and latitude in the form ofcovari- 192 in phenotype 2). This is a relatively large difference ates demonstrated that geographic coordinates had no when compared to established interspecific differences. added explanatory power (ANCOVA, df=l,9; P=0.9 and For instance, the difference in ventral count between D. df=l,9; P=0.4 respectively). Thus, there is no evidence of formosus on the one hand and D. kopsteini and D. clinal variation, the transition from one phenotype to the cyanochloris on the other hand is 5 and 19 respectively & other apparently being sudden rather than gradual. (Vogel Van Rooijen 2007). The difference between D. haasi andD.pictus,D.pictus andamanensis,D. humayu- The sudden phenetic transition represents strong evidence ni, D. tristis is 3, 20, 10 and 14 respectively (Van Rooi- & for lineage separation. Thus, the two distinguished phe- jen Vogel, 2008a). Finally, the difference between D. notypes are here interpreted as representing distinct evo- chairecacos onthe one handandD. tristis andD. schokari lutionary sister-lineages. An evaluation ofthe characters on the other is 10 and 12 respectively (Van Rooijen &Vo- separating the two phenotypes supports this view. First, gel 2009). The third difference between the two pheno- one phenotype exhibits an undivided anal shield where- types is the number of anterior temporal scales. Pheno- as the other has a divided anal shield. An undivided anal type 1 usually has two anterior temporal scales whereas shield is an exceptional character in the genus Den- phenotype 2 has a single anterior temporal in the major- drelaphis. The only congeneric species that exhibits this ity ofcases. This character alone already leads to a cor- character (in roughly 40% of the specimens) is Den- rect identification in 85% ofthe cases. Table 2. Comparison between the type ofDendrelaphis biloreatus and six specimens ofthe species so far referred to as Den- drelaphis gorei. type Dendrelaphis biloreatus Dendrelaphis gorei (n=6) Ventrals 192 190-199 Subcaudals 147 144-152 Relative tail-length 0.34 0.34-0.35 Supralabials 9 8-9 Supralabials touching the eye 3 2-3 Anterior temporals 1-2 1 Total number oftemporals 8 7-12 Poctoculars 2 1-2 Loreals 2 1 Dorsal rows at midbody 13 13 Divided anal shield yes yes Faint ventrolateral line, not bordered by black lines yes yes Postocular stripe that covers the whole yes yes temporal region and extends onto the neck Total length 70.0 53.5-90.0 vertebrals strongly enlarged yes yes Ground color brown yes yes Maxillary teeth 21 20 (n=l) Bonn zoological Bulletin 60 (1): 17-24 ©ZFMK A new Dendrelaphis from Myanmar 21 Fig. 2. Dendrelaphis walli sp. n., holotype (CAS 234880). Nomenclature Two available names, Dendrophis gorei Wall, 1910 and the morphological characteristics ofthe type ofD. bilore- Dendrelaphis biloreatus Wall, 1908 are relevant in the atus and six specimens ofD. gorei. With the exception of context ofthis study. In figure 1, data pertaining to a syn- the loreal shield, the values pertaining to the type ofD. type ofD. gorei and the type ofD. biloreatus are indicat- biloreatus all fall within the range ofD. gorei. Thus, the ed. It is evident thatneitherofthese types represents phe- results agree with Smith'sviewthat the type ofD. bilore- notype 1 (South Myanmar and neighbouring mountain atus represents an exceptional specimen with a split lo- chains). Thus, we here describe phenotype 1 as a new real shield withinD. gorei. In the context ofa revision of species. Phenotype 2 (Assam and Northwest Myanmar) the taxonomy ofthis genus, the authors have come across is currently known as D. gorei (Wall, 1910), but we here several specimens ofDendrelaphis with either a double adoptthe oldernameDendrelaphis biloreatusWall, 1908 or an absent loreal shield. Furthermore D. biloreatus has to representthis lineage. Dendrelaphis biloreatus was de- arather long and slenderpostnasal scale which looks like scribed by Wall on the basis ofa single specimen. Smith second loreal shield at the first glance. So indirectly the (1943) synonymized D. biloreatus with D. gorei as he namemakes some sense. Consequently, the two names in- could not find any difference between the two species deed refer to the same species. According to the rule of apart from the divided loreal shield in D. biloreatus. In priority (ICZN, 1999, art. 23.1), the name Dendrelaphis table 2, a more detailed comparison is provided between biloreatus should actually be adopted for this species. Bonn zoological Bulletin 60 (1): 17-24 ©ZFMK 5 22 Gemot Vogel & Johan van Rooijen Table 3. Morphological and coloration characters ofthe types ofDendrelaphis walli sp. n. Collection N° CAS 234880 CAS 208429 CAS 211939 CAS 222114 CAS 222339 CAS 222 status holotype paratype paratype paratype paratype paratype cSex Ic m m m Ir Snout-vent length (cm) 4A8O.5C 4A9A.A0 5C4A.5C 61.5 5e9n.5c Tail-length (cm) 22.3 25.5 29.5 o29a.0a j">11.A0 Head-length (mm) 1j.j 1J.J 1j.j 1 1.0 t11/.Au Eye-diameter (mm) J.1 J. 1j.j^ j.v J.J — Snout-width 2.8 3.2 3.4 3.7 3.3 Ventrals 213 211 203 210 212 212 Subcaudals 145 - 148 163 147 159 Dorsal formula 13-13-11 - 13-13-11 13-13-11 13-13-9 13-13-11 Temporal formula 22212/2122 222/222 222/212 212/112 2212/222 222/212 Supralabials 8/8 8/8 8/8 8/8 8/8 8/8 Supralabials touching the eye 4,5/4,5 4,5/4,5 4,5/4,5 4,5/4,5 4,5/4,5 4,5/4,5 Tnfralahials 10/10 10/10 119 10/10 11/10 — Infralabials touched 6,7/6,7 6,7/6,7 6,7/5,6 6,7/6,7 6,7/6,7 by first sublabial Loreals 1/1 1/1 1/1 1/1 1/1 1/1 Postoculars 2/2 3/3 2/2 2/2 2/2 2/2 Number ofscales bordering 4 5 5 6 5 the parietal scales Vertebrals larger than yes yes yes yes yes dorsals ofthe first row Anal shield entire yes yes yes yes yes yes Light ventrolateral stripe yes yes yes yes yes which is faint and is not bordered by black lines Postocular stripe broad, yes yes yes yes yes covering the whole temporal region and extending onto the neck Taxonomy Diagnosis. A species ofDendrelaphis, characterized by the combination of: 1) 13 dorsal scale rows at midbody; Dendrelaphis walli sp. n. (Fig. 2) 2) strongly enlargedvertebral scales; 3) 203-213 ventrals; Dendrelaphis gorei (Wall, 1921) (part.) 4) 145-163 subcaudals; 5) 8 supralabials; 6) supralabials 4 and 5 bordering the eye; 7) 2 anterior temporals in the Holotype. CAS 234880, adult female from Myanmar majority ofspecimens; 8) a short sublabial that touches 2 (Chin State: Mindattownship, MindatDistrict, 21 26 43.5 infralabials; 9) an undivided anal shield; 10) a black pos- N, 94 00 24.0 E, 3582 ft), collected byA.K. Shein and T. tocular stripe that covers the majority ofthe temporal re- Nyo, 23 Aug 2005. gion and extends onto the neck; 11) black, oblique bars on the neck region; 12) a pale ventrolateral line, not bor- Paratypes. CAS 208429, CAS 211939, CAS dered by black lines. 222114-222115, CAS 222339. Bonn zoological Bulletin 60 (1): 17-24 ©ZFMK A new Dendrelaphis from Myanmar 23 Table 4. Diagnostic differences between Dendrelaphis walli sp. n. and Dendrelaphis hiloreatus Wall, 1908. Dendrelaphis walli n. sp. (n=7) Dendrelaphis biloreatus (n=7) Ventrals 211(203-213) 192 (190-199) Anterior temporals usually 2 (92% ofspecimens) usually 1 (86% ofspecimens) anal shield entire divided Description of the holotype. Adult female; body very Variation. Table 3 provides data regarding the types of slender; snout-vent length 48.5 cm; tail-length 22.5 D. walli sp. n. cm;head distinct from neck; head-length 13.5 mm; snout- width 2.8 mm; pupil round; eye-diameter3.1 mm; distance Comparison with congeneric species.Dendrelaphiswal- eye-nostril 3.1 mm; 213 ventrals; 145 subcaudals; dorsal li sp. n. differs from all congeners, except Dendrelaphis scales in 13-13-11 rows; 8 supralabials, 4th and 5th bor- biloreatus,Dendrelaphis ca.udolinea.tus andDendrelaphis derthe eye; 10 infralabials, infralabials 1-5 touch the first caudolineolatus,by its dorsal formula of13-13-11/9. Itdif- chinshield, infralabials 5 and 6 touch the second chin- fers from Dendrelaphis caudolineatus in its strongly en- shield; 1 preocular; 2 postoculars; 1 loreal; temporal for- largedvertebral scales (not enlarged inD. caudolineatus), mula 2:2:2:1:2 (L), 2:1:2:2 (R); first sublabial touches in- its much more slender body (stout in D. caudolineatus), fralabials 6 and 7; vertebrals strongly enlarged, with its undivided anal shield, the numberofsupralabials (9 vs. straight posterior margin, width ofthe vertebral scale at 8 in D. walli sp. n.) and in its coloration. It differs from the position ofthe middle ventral scale 2.4 mm; anal en- Dendrelaphis caudolineolatus in the number of ventral tire; parietal scales bordered posteriorly by 4 scales; scales (149-175 vs. 203-213 inD. walli sp. n.) and sub- ground color brownish; supralabials and throat white; caudal scales (111-129 vs. 145-163 inD. wallisp. n.) and some black spots on supralabials 2—4, the loreal and the itsundividedanal shield. Differences betweenD. walli sp. preocular; a black postocular stripe starts behind the eye, n. and D. biloreatus are given in Table 4. covers the majority of the temporal region, and extends onto the neckwhere itbreaks up into narrow, blackoblique Sexual dimorphism. Females have a wider snout than bars which fade away further posteriorly; a faint ventro- males (ANCOVA, df=l,5, P=0.004). In addition, there is lateral line is present, covering the first dorsal row and some evidence that females have a larger eye than males lowerhalfofthe second dorsal row; the ventrolateral line (ANCOVA, df=l,5, P=0.08). No evidence ofother sexu- is not bordered by black lines; belly whitish. al dimorphisms was found though this maybe due to lack ofstatistical power (e.g. Streiner 1990). Distribution. According to currently known locality records, D. biloreatus inhabits the Assam Valley and ad- joining Himalayan foothills whereasD. walli sp. n. inhab- its the South Myanmar lowlands and neighbouring mountain chains. Thus, the two species appear to exhib- it a disjunct distribution (figure 3). However, this maybe due to collecting gaps. In reality, one or both of these species may inhabit the intermediate area. On the other hand, the Myanmar Herpetological Survey did collect in the central andnorthern parts ofMyanmar, which did not yield a specimen ofeither species, with the exception of oneD. biloreatus (CAS 244037) nearthe borderwithAs- sam, within the known range of this species. If these species indeed occur allopatrically, the central dry zone ofMyanmar and adjoining mountain ranges may be the geographic barrier that separates these species and that may have enabled the independent evolution ofthe two. Etymology. This species is dedicated to MajorFrankWall Fig. 3. Currently known distribution of D. biloreatus (1868-1950), in recognition ofhis outstanding work on (white b) and D. walli sp. n. (black W). the genus Dendrelaphis. Bonn zoological Bulletin 60 (1): 17-24 ©ZFMK 24 Gemot Vogel & Johan van Rooijen Acknowledgements. We thank J. V. Vindum (CAS, San Fran- Van Rooijen J, Vogel G (2008a) A review ofthe Dendrelaphis cisco, USA), C. J. 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