Corresponding authors: V. Deepak (
Academic editor Uwe Fritz
During the past 200 years,
Based on a combination of characters including body pattern, external morphology and osteological differences
Specimens of Banded racer found in the southeastern part of peninsular India are morphologically and genetically distinct from populations of the rest of the distribution area. These populations from central and southern Tamil Nadu state represent a different species, consequently described as a new species herein. Furthermore, examination of specimens of the Banded racer from different populations across its entire range, including the type specimen of the genus
The Banded racer, presently named
Our critical analysis of the original source (
We generated DNA sequence data for eight specimens of
Genomic DNA was extracted from liver or tail tissue samples stored in absolute ethanol at –20°C. DNeasy (Qiagen) blood and tissue kits were used to extract DNA. We amplified partial sequences of two nuclear (
Apart from the five genes for which we generated data, we also included an additional gene
All phylogenetic analyses were carried out using the CIPRES Science Gateway portal v3.3 (
In addition to the type series and referred material (n=11) of the new species described below, comparative material examined directly includes the holotype of
Total length, snout-vent length and tail length were measured with thread and a ruler to the nearest 1 mm. Other dimensions were recorded with dial callipers, to the nearest 0.1 mm. Identification of sex was performed by dissection (presence or absence of hemipenes, inspected through a subcaudal incision at the tail base).
The hemipenis of freshly euthanized specimens were everted by pressing the organ until it was everted to the maximum extent, after which the retractor muscle was cut at the level of the 10th subcaudal scale. From preserved specimens in which the hemipenis was not everted, it was removed through a longitudinal incision at 15th subcaudal and detached from the body by cutting the retractor muscle. The organ was separated by making an incision around its circumference at the cloacal region and was immersed in warm water (60°C) for about 5 minutes to soften the tissue. It was then slowly everted with a blunt-end forceps by gently pushing the organ from the distal to proximal end. Afterwards, the hemipenis was softened by again immersing it in warm water for intervals of 5–10 minutes. While everting, petroleum jelly was applied to the inner edge for smooth sliding of the inner ornamented layer. Care was also taken to avoid puncturing the hemipenial wall due to the rubbing of the spines. After eversion, the organ was inflated with petroleum jelly and tied at the base with a thread. Later it was immersed in 1% alizarin red solution in 70% ethanol and removed from the solution after one hour of staining. The calcified spicules were stained using this process and excess stain was removed by immersion of the hemipenis in 70% alcohol for approximately 30 minutes. Detailed photographs of the stained hemipenes were taken using Nikon D 5100 with a 100 mm macro lens (Nikon). Descriptions of hemipenial characters and terminology follow
Length and width of head scales were measured at the longest and widest points of the respective scale(s). Interocular width was measured on top of the head at the shortest distance between the eyes. The number of ventral scales was counted according to
To evaluate character states of midbody vertebrae we created µCT-scans of postcranial skeletons for six “Banded racers”, four
To obtain counts of teeth by a non-invasive procedure, heads of
The opensource software QGIS (
Both BI and ML analysis recover a well supported monophyletic
Map showing geomorphological features in parts of the Indian subcontinent and current locality records (see Appendix
ML phylogeny showing inferred phylogenetic relationships of Old World racers. Labelled in pink colour are the sequences generated in this study, coloured blue is a sample from this study+GenBank sequence. Numbers at internal branches are ML bootstrap values. Bootstrap values below 50 are not shown.
On the same basis and almost simultaneously as
In his “Catalogue of Colubrine snakes […] of the British Museum”
It is also to be noted that from the initial confusion caused by
In this work, we clarify the identity of
To determine the species affiliation we carefully compared Russell’s description and the sketch on plate 21 with published data of
Comparison of available characters for the type specimen of
|
|
|
|
|
|
|
192 | 62 | no | absent | yellow with dark brown center | short whitish bands stippled with black not extending onto ventral scale edges | |
♂♂ 174–186 ♀♀ 186–197 | ♂♂ 63–64 ♀♀ 59–74 | no | absent | creamy white or yellow with dark brown center | short yellowish or whitish bands stippled with black not extending onto ventral scale edges, or patternless | |
♂♂ 180–206 ♀♀ 191–215 | ♂♂ 65–72 ♀♀ 57–73 | yes | present / yes | whitish | creamy-white bands extending onto ventral scale edges | |
|
198–214 | 66–69 | no | present / yes | light brown with irregular dark brown speckles | irregular dirty whitish bands with dark brown speckles extending onto ventral scale edges |
|
197–220 | 69–94 | scarcely | present / yes | dark greyish-white | dirty whitish-brown bands extending onto ventral scale edges |
|
210–224 | 65–72 | yes | present / usually | yellow | without or with very faint yellow bands not extending onto ventral scale edges |
|
165–183 | 53–63 | no | absent but a small spot present | white | yellow bands extending onto ventral scale edges |
|
167–188 | 52–74 | no | absent | whitish-grey | overall speckled with small yellow spots |
|
163–187 | 60–76 | yes | present /yes | whitish | yellow-white bands extending onto ventral scale edges |
|
176–206 | 64–76 | yes | present /scarcely | greyish, mottled with brown | yellow bands extending onto ventral scale edges |
|
154–195 | 35–50 | no | present / yes | whitish | whitish-brown or yellowish bands extending onto ventral scale edges |
|
179–207 | 45–75 | yes | absent | whitish-brown | short white bands not extending onto ventral scale edges |
In a recent study on population systematics of the
Our comparison of the original information of
Based on these consistencies of its morphological characters, colour and pattern, we consider
With the reallocation of the taxon
According to
Some old records mention a specimen under the name
Recently,
It is undisputed that Blyth (l.c.) considered the resemblance of
The preceding list of synonyms and chresonyms contains quotations which to our best knowledge refer to the snake species so far known as Banded racer (
Extract from a previously unpublished original water-colour painting from Brian H. Hodgson’s collection of drawings kept in the Zoological Library of the Natural History Museum, London, U.K. The depicted specimen from the „Central hilly region “of Nepal, determined by
Franz
We also tracked down two older and previously overlooked or wrongly assigned names that clearly refer to what is currently understood to be the Banded racer, viz.
From the mid of the 19th century onwards both names fell into oblivion as potential synonyms of the Banded racer. This was probably also due to
To our best knowledge,
We could not locate the two type specimens of
The same applies to the type specimen of
Based on our own data and summarized information from literature (
Pholidosis: Head 2.17–2.14 times longer than broad (males 1.17–2.14, females 1.22–2.05), canthus rostralis moderate developed. Rostral nearly twice as broad as high, clearly visible from above. Internasals usually shorter than prefrontals, sometimes equal in length in, e.g.
Predominantly eight supralabials, last two or three being larger (longer), fifth highest; nine in
Dorsal scale reduction formula summarized from 31 examined specimens (see Appendix
2+3, 3+4, or 10+11 (6–11) 2+3, 3+4, 7+8, 8+9, 9+10, or 10+11 (65–131) [79–133]
3+4, or 10+11 (5–9) 2+3, 3+4, 7+8, 8+9, 9+10, or 10+11 (63–131) [82–129]
2+3, 3+4, 4+5, 8+9, 9+10, or 10+11 (68–145) [105–137]
1+2, 2+3, 3+4, 4+5, 7+8, 8+9, or 9+10 (68–142) [107–138]
4+5, 6+7, 7+8, 8+9, or 9+10 (111–192) [157–174] 7+8 (188–194)
3+4, 4+5, 7+8, 8+9, or 9+10 (112–191) [156–178] 7+8, or 8+9 (190–195)
Six examined specimens (
The hemipenis reaches up to 13th subcaudals and is 36.9 mm in length and 8.2 mm maximum width at midbody and apex. It is divided into three distinct areas; the proximal area (nearly 30% of the total hemipenial length) is smooth without any ornamentation, middle zone (about 25% of the total hemipenial length) is with evenly scattered spinules and the apical calyculated portion (45% of the total hemipenial length). No enlarged spines are present, but the size of spinules at the proximal end are slightly larger than the ones at the distal end. The calyculated area can be further divided almost equally into proximal half with large calyces and distal half with smaller and denser calyces. The sulcus spermaticus is single, bounded with thick walls, runs straight across its length and opens into a delta (3 mm wide at apex, 1.9 mm in length), which is nude. The delta is subapical in position and the apex is calyculated, edged with fewer papillae. At the sulcate side, the calyces are larger proximally and gradually smaller and denser towards the distal end. The distal calyces are scalloped and edged with papillae. Along the asulcate side the spinous area starts more distally than the sulcate and lateral side. After the spinous zone, there are 2–3 rows of large calyces followed by densely packed calyces towards the apex. The proximal large calyces (9–10 in number) are of uneven size (mostly rounded or pentagonal) and among them the distal ones are scalloped. The large scalloped calyces are of 2.2–2.7 mm in length and 1.9–2.5 mm in width.
15/15 maxillary teeth, the anterior 13/13 precranterian teeth are increasing in size posteriorly and are followed by a small diastema and 2/2 enlarged roundish cranterian teeth without groove. All maxillary teeth are slightly curved posteriorly, without significant interspaces. Medial to the precranterian teeth are single replacement tooth at different growth stages. One to two replacement teeth per cranterian tooth are found posteromedially to each tooth, showing different growth stages. 11/11 palatine teeth, decreasing in size posteriorly. All are curved posteriorly shortly above the base. Lateral to the palatine teeth there is a single replacement tooth at different growth stages. No significant interspace exists between the different palatine teeth. Posteromedial process of palatine large, expanding two pterygoid teeth. 13/12 pterygoid teeth, decreasing in size posteriorly. All are curved posteriorly shortly above the base. Lateral to the pterygoid teeth there is a single replacement tooth at different growth stages. No significant interspace exists between the different palatine teeth. The posterior 57% of the pterygoid are without teeth. 17/17 mandibular teeth, increasing in size up to tooth 5/5 and from there decreasing in size posteriorly. All are slightly curved posteriorly. Medial to the mandibular teeth there are 1–2 replacement teeth at different growth stages.
Four specimens from Pakistan (
The body is robust, moderately stout, roundish in cross-section at midbody. Head moderately pointed, barely distinct from neck. Eye large, 16–24% of head length. Male specimens grow slightly larger than females; the longest examined specimens are from Sind, Pakistan including a male (
Dorsal colour and pattern show a significant ontogenetic change. In juvenile and subadult specimens the dorsal ground colour is olive-brown to light brown, the head dorsally slightly darker with narrow elongated irregular shaped whitish markings. Along the body up to 70 narrow whitish/black or dark brown stippled crossbars, usually two scales wide and in contact with lateral edges of ventrals. The first band is separated from the posterior border of parietals, the interspace between bands is three to four dorsal scales wide. The intensity of bands decreases posteriorly and the contours become blurred near to the vent. Dorsocaudally the bands are dissolved in an irregular fine pattern of light and dark points. Temporals, preoculars and supralabials can show small whitish spots. From about 600 mm total length the colour and pattern of young specimens begin to fade and disappear gradually. Adult specimens are dorsally dark brown to dark reddish-brown without any markings on head, body or tail. The ventral side of head, body and tail in all age classes is patternless yellowish cream.
According to our current knowledge,
Hemipenis of
In Pakistan confirmed records are known from the Indus plain in Sindh province (nr. Badin, Hala, Jati, Lakarna, Makli Hills, Mohenjo-daro, Pir Patho, Raj Malk, Sonda, and Tatta) and some authors mentioned southern and central Punjab without specific locality (e.g.
Sporadically the Banded racer is mentioned for the Himalayan region in northwestern and northeastern India.
Based on water-colour sketch no. 61 from the Hardwicke collection,
When it comes to the distribution of the Banded racer in Sri Lanka, there is a consistent reference to
In several publications, especially in overview works and faunal lists, the Banded racer is mentioned for Bangladesh. This is probably because in older publications (e.g.
From Nepal, the Banded racer was first mentioned by
Based on records of the Banded racer in Bengal,
Skull of
Habitats of
The vertical distribution ranges from sea level to over 1000 m altitudes, with about 85% of all finds below 600 m a.s.l. A few unusually high sites are known from the states of Madhya Pradesh (Pachmarhi Biosphere Reserve, 1053 m a.s.l.) and Maharashtra (Kaas Plateau, 1191 m a.s.l., and Panchgani, 1258 m a.s.l.) (see Appendix
It is a mainly terrestrial species but also observed to be a good climber (
It is a diurnal species that is only exceptionally observed active during the night and known for its defensive and pugnacious behaviour. If threatened or captured it erects itself, flattens the forebody and furiously attacks when further provoked which one can confuse with a cobra (
The Banded racer is an active hunter that usually kills its prey by constriction or body pressure. Its prey consists of small mammals like rats, mice, shrews and bats, but also frogs, lizards, birds, worms, and for juveniles insects are part of the food spectrum (
Mating of the Banded racer was observed between late winter and mid-monsoon (
Manhas (2015a) observed the Banded racer as roadkill during monsoon in Bhopal district of Madhya Pradesh state and
For Pakistani specimens,
A medium sized (maximum total length 951 mm) snake with countersunk lower jaw; dark brownish dorsum; head with irregular white spots, two slanting roughly “∏” shaped white markings with black edges on either side, starting on the back of the head (behind parietals) extending into the body, almost the length of head; 13–18 prominent white bands in the anterior region of the dorsum in both juveniles and adults; 34–48 total bands on the body in both juveniles and adults; 23:21(exceptionally 23):16–18 smooth dorsal scale rows; 189–218 ventrals (males: 192–197; females: 189–216); 76–88 subcaudals (males: 83–88; females: 76–88); cloacal plate divided; tail without bands and underside creamish. Its dentition is characterized by 12–16 maxillary teeth, the last two enlarged and separated by a diastema, 9–11 palatine teeth, 14–15 pterygoid teeth and 15–16 mandibular teeth.
NCBS AU-732, adult female, from Tuticorin, Tamil Nadu state, India (
Holotype of
Line drawings showing head scalation and dimensions of the holotype of
The specific epithet is a patronym of late Mr Naveen Joseph. Naveen was a naturalist from Tuticorin, well known for his research on reptiles, particularly snakes in that region. He was a friend of VD, SN, and GM and helped them collect specimens of the new species.
Morphometric and merestic data are provided in Table
Meristic and morphometric (in mm) data for examined specimens of
|
|
|
|
|
|
|
|
|
|
Sex | female | male | female | female | male | female | female | female | unknown |
|
757.6 | 574.3 | 592.0 | 608.5 | 440.0 | 255.0 | 574.5 | 722.2 | NA |
|
175.5 | 167.1 | 162.2 | 89.9 | 142.0 | 60.0 | 170.5 | 228.9 | NA |
|
933.1 | 741.5 | 754.2 | 698.4 | 582.0 | 315.0 | 745.0 | 951.1 | NA |
|
12.4 | 11.9 | 10.8 | 13.5 | 11.6 | 7.9 | 13.4 | NA | NA |
|
24.1 | 20.0 | 21.6 | 22.7 | 14.5 | 10.7 | 23.0 | NA | NA |
|
4.1 | 3.4 | 3.9 | 3.8 | 3.3 | 2.7 | 3.7 | NA | NA |
|
4.8 | 4.8 | 4.2 | 4.6 | 3.8 | 2.4 | 4.1 | NA | NA |
|
6.4 | 6.1 | 5.6 | 5.3 | 5.2 | 4.3 | 5.9 | NA | NA |
|
4.7 | 3.8 | 4.3 | 3.7 | 3.7 | 3.0 | 4.2 | NA | NA |
|
2.0 | 2.3 | 2.2 | 2.2 | 2.2 | 1.6 | 2.0 | NA | NA |
|
1.2 | 1.6 | 1.6 | 1.5 | 2.1 | 1.5 | 1.9 | NA | NA |
|
8.2 | 7.6 | 7.2 | 7.7 | 6.3 | 4.6 | 7.3 | NA | NA |
|
6.3 | 5.7 | 6.0 | 6.0 | 5.7 | 4.5 | 5.8 | NA | NA |
|
4.5 | 4.2 | 4.2 | 4.1 | 3.9 | 3.0 | 4.6 | NA | NA |
|
23/21/18 | 24/23/17 | 23/21/17 | 23/21/17 | 21/21/17 | 23/23/17 | NA | 23/21/16 | NA |
|
3 | 1 | 2 | 0 | 2 | 1 | 3 | 2 | NA |
|
203 | 192 | 195 | 208 | 197 | 216 | 204 | 189 | 218 |
Subc | 76 | 82 | 77 | 84 | 88 | 79 | damaged | 88 | NA |
|
283 | 276 | 275 | 293 | 288 | 297 | NA | 280 | NA |
|
2/2 | 2/2 | 2/2 | 2/2 | 2/2 | 2/2 | 2/2 | 2/2 | 2/2 |
|
2/2 | 2/2 | 2/2 | 2/2 | 2/3 | 2/2 | 2/2 | 2/2 | 2/2 |
T | 2+2/2+3 | 2+3+4/2+2+4 | 2+3/2+3 | 2+3/2+3 | 2+2+3/2+3+4 | 2+3+4/2+2+4 | 2+3+4/2+2+4 | 2+3/damaged | 2+3/2+3 |
|
8/8 | 8/8 | 8/8 | 8/8 | 8/8 | 8/8 | 8/8 | 8/8 | 8/8 |
|
9/9 | 10/10 | 10/10 | 10/10 | 11/11 | 10/10 | 10/10 | damaged | 10/10 |
Body subcylindrical, dorsoventrally flattened. Head ovate, barely wider than the anterior end of body. Total lenghth 934 mm; snout-vent length 758 mm; tail length 176 mm; ratio tail length/total length 0.19; head length 24.0 mm; rostral large, 2.2 times broader than high, not protruding, rounded in dorsal view, and wedged in between internasals; the latter smaller (1.5 mm) and distinctly shorter along median suture than prefrontals (2.5 mm); mid-line suture between internasals and pre-frontals in straight line; distance from posterior tip of rostal to anterior edge of frontal 4.2 mm, the latter bell-shaped, 6.4 mm long with a maximum width of 4.7 mm; interocular width 7.0 mm; parietals 6.3 mm long, outer lateral and posterior margins of parietals surrounded by 13 scales between the upper postoculars; nasal completely divided, in contact with first and anterior half of second supralabial; nostril large in anterior shield; distance between posterior margin of nostril and anterior border eye 4.2 mm; loreal slightly longer (1.5 mm) than high (1.4 mm), in contact with posterior part of second and anterior half of third supralabial; preocular single, reaching top of head, touching frontal; presubocular squarish, smaller than preocular and loreal, in contact with third and fourth supralabial; 8/8 supralabials, fourth and fifth entering the eye, fifth highest; 2/2 postoculars, upper slightly broader, lower higher, lower postocular in contact with fifth supralabial; temporals in 2+3/2+3 rows, lower anterior slightly larger than upper; 9/9 sublabials, sixth largest, first five in contact with anterior inframaxillaries on both sides; anterior pair of inframaxillars broader and slightly longer (6.1 mm) than posterior pair (5.6 mm), which is distinctly separated by five inserted gulars; five gulars in a row between posterior edge of anterior inframaxillars and preventral; three preventrals and 203 ventrals; anal scute divided, right overlapping left; 76 paired subcaudals; terminal spine blunt; dorsal scales smooth, arranged in 23/21/17; apical pits present, but not consistently on all the dorsal scales, single or paired on dorsal part from the neck to near sacral region, single or triple pits present on dorsocaudal scales.
Dorsal scale reduction formula:
-3(6) +10(64) -10(67) +10(82) 10+11(87) 3+4(133) 8+9(151) –
-4(8) +11(65) -11(69) +11(84) -11(88) 3+4 (133) 8+9(150) +2(202)
14/16 maxillary teeth, the anterior 12/14 precranterian teeth are increasing in size posteriorly (from 0.9 to 1.6 mm length) and are followed by a diastema which is as long as the socket of the last precranterian tooth, and 2/2 slightly enlarged roundish cranterian teeth (~1.9 mm long) without groove. All maxillary teeth are slightly curved posteriorly, without significant interspaces. Medial to the precranterian teeth are single replacement tooth at different growth stages. One to 2 replacement teeth per cranterian tooth are found posteromedially to each tooth, showing different growth stages.
9/9 palatine teeth, decreasing in size posteriorly. All are curved posteriorly shortly above the base. Lateral to the palatine teeth there are single replacement tooth at different growth stages. No significant interspace exists between the different palatine teeth. Posteromedial process of palatine large, expanding two pterygoid teeth.
14/14 pterygoid teeth, decreasing in size posteriorly. All are curved posteriorly shortly above the base. Lateral to the pterygoid teeth there are single replacement tooth at different growth stages. No significant interspace exists between the different palatine teeth. The posterior 56% of the pterygoid are without teeth.
15/15 mandibular teeth, increasing in size up to tooth 6/6 and from there decreasing in size posteriorly. All are slightly curved posteriorly. Medial to the mandibular teeth there are 1–2 replacement teeth at different growth stages.
The hemipenis reaches up to 12th subcaudals and is 24.3 mm in length with a maximum width of 5.8 mm at 1/3rd of the proximal end. It is divided into three distinct areas; the proximal area (nearly 1/4th of the total hemipenial length) is smooth without any ornamentation, middle zone (about 28% of the total hemipenial length) is with spinulae and the apical calyculated portion (48% of the total hemipenial length). The spicules at the midbody of the hemipenis gradually reduce in size from proximal to the distal end. The calyculated area can be further divided almost equally into proximal half with large calyces and distal half with smaller and denser calyces. The sulcus spermaticus is single and runs straight across its length and opens into a wide and elongate naked area (3.3 mm) extending laterally from the proximal to distal end extending to the apex. The opening of the sulcus forms a triangular area and ends apically. The sulcus is bounded with thick walls. The calyces are larger proximally and gradually smaller and denser towards the distal end. The calyces distally are edged with papillae. Along the asulcate side, the spinous area starts more distally than the sulcate and lateral side. The proximal calyces are wider and elongated, with eight to nine well defined calyces extending towards the lateral side and the distal part of these calyces are scalloped. The calyces towards the distal part are with more of papiliated edges and are similar in micro-ornamentation with the sulcate side.
Hemipenis of
Skull of the holotype of
See Table
NCBS-AU733 differs from the holotype in having a slightly darker frontal and suprocular scales, presence of irregular white spots in the temporal scales, and a small white line along the midline between the two prefrontals. The first band on the body is separated from the “∏” shaped markings by 2 scales on both the sides along the lateral side. 46–48 bands on the dorsum from neck to vent, anterior 18 bands are prominent with white and black edges which fades and become wider towards the vent.
The male paratype from “Salem” (
Head 1.25–1.99 times longer than broad (male 1.25–1.67, females 1.35–1.99). Rostral twice as broad as high and visible from above. Internasals (in midline suture) usually smaller than prefrontals. Frontals 1.32–1.59 times longer than the maximum width (male 1.40–1.59, females 1.32–1.42). Posterior border of parietals uneven in shape at the median suture forming an acute angle in
Dorsal scale rows (
Dorsal scales with single or paired apical pits. Supracaudal scales usually with one to three pits on the first three to five scales followed the supracaudal reduction. In male (
Dorsal scale reduction formula summarized from five examined specimens (see Appendix
4+5(6) 3+4(6–13) 2+3 or 3+4(116–133) 7+8 or 8+9 (144–151)
– 2+3 or 3+4(6–13) 3+4 (115–133) 7+8 or 8+9 (142–151)
Two examined specimens (NCBS-AU732, holotype, and NCBS-AU733) and one specimen (
The body robust, moderately stout, roundish in cross section at midbody. Head moderately pointed, barely distinct from neck or sometimes slightly distinct. Eye large with black round pupil, 16–25% of head length. The longest examined specimens are from Karur (
Habitats of
The diet of the
Although local envenoming by congeners is reported for
Juvenile
The first detailed description of osteological features of
In his comparison,
Some of the osteological characters used by
Recently,
In contrast to the former studies on
We found differences within the distal ends of the lateral processes of the premaxilla, which either are tapering into a single tip in
Furthermore,
Wilson (l. c.) mentioned that in
Another character listed by Wilson (l. c.) as typical for
Wilson (l. c.) also described the pterygoid processes at the basioccipital which he mentioned as clearly visible in e.g.
Differences in midbody vertebrae morphology of racer species is highlighted as a character of phylogenetic significance (
According to
In summary, it is evident that
1a. | Midbody dorsal scales arranged in 23 rows |
|
1b. | Midbody dorsal scales arranged in 21, exceptionally 23 rows |
|
1c. | Midbody dorsal scales arranged in 19 rows |
|
2a. | Large brownish dark-edged roundish, oval or saddle-shaped dorsal markings extending down both flanks, 118– 27 subcaudals |
|
2b. | Two clearly demarcated slanting rougly “∏” shaped white markings on the back of head, 76–88 subcaudals |
|
3a. | Large brownish dark-edged oval or saddle-shaped dorsal markings extending down both flanks (nape element may be elongated and confined to middorsal segment, SE India) |
|
3b. | Dorsal colour pattern not as described (from western Nepal [unconfirmed] and northern India westward) |
|
4a. | With a reddish vertebral stripe |
|
4b. | Vertebral stripe absent, postsubocular absent (two supralabials entering eye) or present (oftenso in |
|
5a. | Fewer than 220 ventrals and 332 ventrals plus subcaudals |
|
5b. | More than 219 ventrals and 331 ventrals plus subcaudals |
|
6a. | Nuchal streak present (very rarely absent), dorsal colour pattern along entire trunk |
|
6b. | Nuchal streak absent (very rarely present), last quarter of trunk devoid of pattern or unicoloured throughout |
|
7a. | 199 – 204 ventrals, 108 to ca. 112 subcaudals, sum 307 to ca. 314 (only ♂♂ known, probably close to and above 1,800 m a.s.l. east of Nushki, Urak Valley area, and Ziarat Mountains in NE Balochistan, Pakistan) |
|
7b. | More than 203 ventrals and 105 subcaudals, sum at least 318 (minima from Hindukush to N Khyber Pakhtunkhwa) |
|
8a. | 110–127 subcaudals, postsubocular present or absent (0 – 2 supralabials entering eye), preocular often divided (northern Balochistan Region) |
|
8b. | 106–148 subcaudals (132–144 in |
|
9a. | 230–247 ventrals, 133–148 subcaudals, sum 363–395 (documented for Sindh and contiguous Balochistan Province, probably also SE Punjab and possibly W Rajasthan) |
|
9b. | 220–256 ventrals (fewer than 220 in area of potential sympatry with |
|
10a. | Usually fewer than 233 ventrals (as much as 244 in Chagai), 19 dorsal scale rows on anterior trunk or at midbody and 11–13 prior to vent |
|
10b. | 237–256 ventrals, 19–21 dorsal scale rows on anterior trunk or at midbody and 13–15 prior to vent (eastern Hindu Kush, Karakoram, Ladakh, Spiti Valley in NE Himachal Pradesh, and probably Zanskar) |
|
The maximum likelihood (ML) and Bayesian (BI) phylogeny presented in this study were similar but not same, there were some unresolved nodes in the BI phylogeny. Furthermore, the phylogenetic relationships are different from that presented by
In their study on Western Palearctic colubrid snakes,
We thank Sameresh Changdar, Vikash Behari and Parag Nag at GE Inspections Technologies, Chakan, Pune for their support and help in microCT scanning of the holotype of Joseph’s racer. We would like to thank Anil Khaire (Chairman Indian Herpetological Society, Pune), Vishal Verma, Sandeep Belkhede, Madhao Vaidya and Varad Giri (NCBS, Bangalore), V.J. Jins (SACON, Coimbatore), Chethan Kumar (Wildlife Institute of India, Dehradun) Raju Vyas (Gujarat) and Caleb Daniel (IISc, Bangalore), Suvendu Swain and Rakesh K. Mohalik (Odisha) for donating tissue samples for this study. We would like to thank Praveen Karanth for the lab facilities, Chinta Sidharthan and Kunal Arekar for support during lab work and Natalie Cooper for help with some of the analyses. We would also like to thank S.P. Vijayakumar for his help and inputs on microCT scanning. We are grateful to Naveen Joseph’s family and friends (Akila Joseph, Eswarapandi, Mahendraraja, Zakaria) for their support.
We are indebted to curators and other staff who approved loans made us welcome at their collections or provided photographs and information about specimens under their care used for this study. These are Rahul Khot, Saunak Pal, Mrugank Prabhu, Vithoba Hegde, Parvathy and Shyam Jadhav (
We are grateful to Rea Kavazi and Gillian Humphreys (Bodleian Library, Oxford, U.K.), David J. Gower (NHM London U.K.), John Rose, Andrea Hart and Rosie Jones (NHM Library, London, U.K.) who assisted us in the search for unpublished manuscripts and original drawings from T. E. Cantor, B. H. Hodgson and T. Hardwicke, and permitted us to reproduce them. For the support in procuring rare literature, we thank Hans-Ulrich Raake and Martina Rißberger (Zoological Main Library, Museum für Naturkunde Berlin, Germany), as well as Bernhard Schurian (Museum für Naturkunde Berlin, Germany) for scanning and postprocessing of the plates, reproduced from Russell`s book. Christoph Kucharzewski (
Francis Gowdie (date of birth unknown –12. Sept. 1813), served in the Third (1789–92) and Fourth (1798–99) Anglo-Mysore War and from 1805 to 1813 as Major General in the service of the East Indian Company Army at Madras (Chennai, Tamil Nadu state of India) (
Earlier attempts by
But “Caser cotta” lies on the northern part of the Malabar Coast and therefore opposite of the Coast of Coromandel where Russell spend most of the time during his medical service for the East Indian Company. Furthermore,
The description of
We here provide the transcript of the detailed description of
The title page contains following additional text: “A catalogue of the species will appear in the Proceedings of the Zool. Soc. London. April 29. 1839”, and the following information about the content of the manuscript: “35 Drawings {28 Figures of Serpents. New Species., 7. Anatomical details of Do.”, and “ The descriptions occupy 240 pages”.
The description of
“
Above partly bright/ olive green with white/ and|
Dimensions/
ft. in lin
Breadth of the head 0. 0. 5/
Length of the head* 0. 1. 0/
Length of the trunk 2. 4. 6/
Length of the tail 0. 7. 0/
Total length …… 3 ft. 0 in. 6 lin/ Circumference of the neck. 1 inch 3 lines./ Greatest circumf. of the trunk: 2 inches./
*The upper surface of the/ head being strongly arched,/ the measurement is taken on/ the lower surface from the projecting muzzle to the anterior margin/ of the third scutum abdominal |
The alleged collector Sir William Crofton Twynam (1827–1922) also known as “Raja of the North” was born in Galle. He entered the Ceylon Civil Service in 1845 and held from 1869 until his retirement in 1896 the post as Government Agent on Jaffna (
GenBank sequences number and locations for the samples used in this study. “_” no data or information not available. Sequences generated in this study marked in bold.
Species | Locations | Voucher number |
|
|
CMOS | 16S | 12S | RAG1 | Genseq |
---|---|---|---|---|---|---|---|---|---|
|
Niokolo-Koba NP, Senegal | HLMD RA2906 |
|
|
|
|
— | — | nomenclature |
|
Niokolo-Koba NP, Senegal | HLMD RA2906 |
|
|
|
|
— | — | |
|
Iran | ICSTZM.7H.1154 |
|
|
— | — |
|
— | |
|
Greece, Serifos | NHMW KCC1 |
|
|
|
|
|
— | |
|
Turkey, Antalya Province | MVZ 230242 |
|
|
|
|
|
— | |
|
Turkmenistan |
|
|
|
|
|
— | ||
|
Turkey | CS4655/J170 |
|
|
|
|
— | — | |
|
Anavarza, Turkey & Osmaniye, Turkey | CS97 Eb1/J252 |
|
— |
|
|
— | — | |
|
Aras, Armenia | ZISP 27859 |
|
— |
|
|
— | — | |
|
Jordan | CS98 cr/J209 |
|
— |
|
|
— | — | |
|
Syria | CS94 de2/J226 |
|
— |
|
|
— | — | |
|
Kahramanmaras, Turkey | CS4653/J166 |
|
|
|
|
— | — | |
|
Turkey | NMPGV 34464 |
|
|
|
— | — | ||
|
Antakya, Turkey & Osmaniye, Turkey | J168, CS4651/J171 |
|
|
|
|
— | — | |
|
Turkey | CS93li/J249 |
|
— |
|
|
— | — | |
|
Hamadan, Iran | MNHG 2627.004 |
|
— |
|
|
|
— | |
|
Sivas, Turkey & W. Turkey | CS4664/J173 |
|
|
|
|
|
— | |
|
Ufra, Halfeti, Turkey | CS00pe/J200 |
|
— |
|
|
|
— | |
|
Turkey | J175 |
|
|
|
|
— | — | |
|
Turkey | CS97 ro1/J225 |
|
— |
|
|
— | — | |
|
Van, Turkey | CS 01th/J194 |
|
— |
|
|
— | — | |
|
Ogooue, Gaboon | — | — | — |
|
|
|
— | |
|
Socotra, Yemen | HLMD RA2973 |
|
|
|
|
|
— | |
|
Morocco | HLMD RA-1187 |
|
|
|
|
|
— | |
|
Cadiz Province, Spain & Morroco | MNCN 11988 |
|
|
|
|
|
|
|
|
Kotajsk Region, Armenia | ZISP 27709 |
|
|
|
|
|
— | |
|
Kotajsk Region, Armenia | ZISP 27733 |
|
|
|
— |
|
— | |
|
Cyprus | Hcyp43 |
|
— | — | — |
|
— | |
|
Narina, Croatia | — |
|
|
|
|
|
— | |
|
Spain | MVZ 178418 |
|
|
|
|
|
|
|
|
Djébil NP, Tunisia | HLMD RA2333 |
|
— |
|
|
|
— | |
|
Khuzestan Province, Iran |
|
|
— | — | — |
|
— | |
|
Pakistan |
|
|
— |
|
|
|
— | |
|
Sam, Rajasthan, India | — | — |
|
— |
|
— | — | genseq4- |
|
Morocco | E2208.8 |
|
— | — |
|
|
— | |
|
Portugal |
|
|
— |
|
|
|
|
|
|
Te-touan, Tleta Tarhremt, Morocco | MVZ 186239 |
|
|
— | — | — | — | |
|
Bou Hedma, Tunisia | HLMD RA2974 |
|
— |
|
|
|
— | |
|
Spain | MVZ 186073 |
|
|
|
— | — | — | |
|
Mallorca, Baleanc I., Spain | E2208.6 |
|
— | — |
|
|
— | |
|
Namibia |
|
|
|
|
— | — | ||
|
Turkey | ZMHRU2014/60-5 |
|
— |
|
|
|
— | |
|
Yunnan, China | MVZ 211019 |
|
|
|
|
|
— | |
|
Iran | RAP-2019 |
|
|
|
— |
|
— | |
|
Turkmenistan |
|
|
|
— |
|
— | ||
|
Hyderabad, India | — | — |
|
— | — | — | — | genseq4- |
|
Jordan | HLMD J14 |
|
|
|
— |
|
_ | |
|
Bangalore, India | NCBS AQ-492 |
|
|
|
|
— | — | genseq4- |
|
Baripada, India | — |
|
— | — |
|
— | genseq4- |
|
|
Choudwar, India | — |
|
|
|
— | — | genseq4- |
|
|
Yavatmal, India | — |
|
— | — | — | — | genseq4- |
|
|
Pune, India | — |
|
— | — | — | — | geneseq4- |
|
|
Egypt | — |
|
|
|
— |
|
— | |
|
Gujarat, India | — |
|
|
|
— | — | — | genseq4- CTYB, |
|
Anaikatti, India | — |
|
— |
|
— | — | genseq3- |
|
|
Tuticorin, India | NCBS-AU732 |
|
|
|
— | — | — | genseq1- |
|
Karur, India |
|
— | — |
|
— |
|
genseq2- |
|
|
Krasnovodsk, Turkmenistan & Uzbekistan |
|
|
|
|
|
— | ||
|
Ahal Prov., Turkmenistan & Ambouli, Djibouti |
|
|
|
— | — | — | ||
|
Sinai, Egypt |
|
|
|
|
|
— | ||
|
Hurkania, Israel | SMNH<ISR>:17853 |
|
— | — | — |
|
— | |
|
Sam, Rajasthan, India | — | — |
|
|
— | — | — | genseq4- |
|
Oman | CN4780 |
|
— |
|
|
|
— | |
|
Israel | TAU. |
|
— |
|
|
|
— | |
|
Ilam, Abdanan, Iran | — |
|
— |
|
— | — | — | |
|
Israel | TAU. |
|
— |
|
|
|
— | |
|
Iran |
|
|
— |
|
|
|
— | |
|
Turkey | ZMHRU2015/0 |
|
— |
|
|
|
— | |
|
Tanzania | CMRK284 |
|
|
|
— | — | — | |
|
Sri Lanka | RAP0491 |
|
|
|
|
|
— | |
|
Egypt | — |
|
— |
|
— |
|
— | |
|
Jordan & Hail, Saudi Arabia | HLMD J62 |
|
|
|
|
|
— | |
|
Gujarat, India | NCBS HA-105 | — |
|
|
|
|
— | |
|
Gujarat, India | — |
|
|
|
|
— |
|
genseq4- |
|
Unknown | — |
|
|
|
— | — | — | |
|
Maharashtra, India | — |
|
|
|
|
— |
|
genseq4- |
Partitions and models of sequence evolution used in the Maximum Likelihood (ML) and Bayesian Inference (BI) phylogenetic analyses. In the second column, 1st, 2nd and 3rd refer to codon position.
|
|
|
|
P1 | GTR+G | GTR+I+G | |
P2 | GTR+G | HKY+I+G | |
P3 | GTR+G | GTR+I+G | |
P4 | GTR+G | HKY+G | |
P5 | GTR+G | HKY+G | |
P6 |
|
GTR+G | GTR+I+G |
Pairwise uncorrected genetic distance matrices showing within and between generic distances for the three mitochondrial genes. Within genus pairwise distances,
|
|||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Genus | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 |
|
|
||||||||||||||||
|
0.18–0.19 | ||||||||||||||||
|
0.14–0.18 | 0.16–0.18 |
|
||||||||||||||
|
0.15–0.18 | 0.16–0.17 | 0.12–0.16 |
|
|||||||||||||
|
0.18 | 0.17 | 0.14–0.17 | 0.14–0.17 | |||||||||||||
|
0.14–0.17 | 0.15–0.17 | 0.12–0.17 | 0.13–0.17 | 0.16–0.18 |
|
|||||||||||
|
0.15–0.17 | 0.17–0.18 | 0.12–0.15 | 0.10–0.15 | 0.15–0.16 | 0.13–0.15 |
|
||||||||||
|
0.16–0.19 | 0.16–0.18 | 0.15–0.17 | 0.15–0.18 | 0.17 | 0.15–0.17 | 0.15–0.16 |
|
|||||||||
|
0.17–0.19 | 0.12–0.15 | 0.13–0.18 | 0.14–0.19 | 0.15–0.17 | 0.15–0.19 | 0.15–0.18 | 0.14–0.18 |
|
||||||||
|
0.19–0.21 | 0.20 | 0.18–0.19 | 0.17–0.20 | 0.19 | 0.18–0.19 | 0.19–0.20 | 0.18–0.20 | 0.15–0.18 | ||||||||
|
0.17–0.19 | 0.18 | 0.15–0.19 | 0.16–0.19 | 0.19 | 0.17–0.18 | 0.17–0.18 | 0.17–0.20 | 0.17–0.21 | 0.21 | |||||||
|
0.17 | 0.16 | 0.13–0.16 | 0.12–0.15 | 0.15 | 0.14–0.16 | 0.12–0.13 | 0.16–0.18 | 0.15–0.17 | 0.18 | 0.18 | ||||||
|
0.16–0.17 | 0.16 | 0.14–0.16 | 0.14–0.16 | 0.16 | 0.15–0.16 | 0.14–0.17 | 0.14–0.16 | 0.15–0.17 | 0.18 | 0.17 | 0.15 | |||||
|
|
0.17–0.18 | 0.13–0.18 | 0.13–0.18 | 0.16–0.17 | 0.13–0.16 | 0.14–0.17 | 0.15–0.19 | 0.14–0.20 | 0.18–0.20 | 0.17–0.19 | 0.15–0.16 | 0.15–0.17 |
|
|||
|
0.17–0.19 | 0.16–0.18 | 0.15–0.18 | 0.15–0.19 | 0.16–0.17 | 0.15–0.18 | 0.15–0.18 | 0.15–0.17 | 0.14–0.19 | 0.20–0.21 | 0.18–0.20 | 0.16–0.17 | 0.14–0.16 | 0.15–0.20 |
|
||
|
0.19–0.21 | 0.19 | 0.17–0.20 | 0.18–0.20 | 0.20 | 0.18–0.20 | 0.19–0.20 | 0.18–0.19 | 0.18–0.20 | 0.20 | 0.20 | 0.20 | 0.18 | 0.19–0.22 | 0.19–0.20 | ||
|
0.15–0.17 | 0.16 | 0.15–0.16 | 0.13–0.16 | 0.16–0.17 | 0.14–0.15 | 0.14–0.16 | 0.16–0.18 | 0.14–0.18 | 0.18 | 0.18 | 0.16 | 0.15–0.16 | 0.14–0.16 | 0.17–0.19 | 0.19 | |
|
0.18–0.20 | 0.19 | 0.16–0.20 | 0.18–0.21 | 0.19–0.20 | 0.17–0.19 | 0.18–0.19 | 0.17–0.19 | 0.19–0.20 | 0.21–0.22 | 0.20–0.22 | 0.18–0.19 | 0.17 | 0.17–0.19 | 0.16–0.18 | 0.19–0.21 | 0.19 |
|
0.15–0.19 | 0.16–0.17 | 0.14–0.16 | 0.14–0.18 | 0.18–0.19 | 0.15–0.18 | 0.13–0.16 | 0.15–0.16 | 0.16–0.18 | 0.18–0.20 | 0.18–0.19 | 0.15–0.17 | 0.16 | 0.14–0.18 | 0.15–0.18 | 0.18–0.19 | 0.16 |
|
|||||||||||||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
||||||||||||||||
|
0.14–0.15 | ||||||||||||||||
|
0.16–0.18 | 0.13–0.15 |
|
||||||||||||||
|
0.14–0.19 | 0.14–0.15 | 0.12–0.16 |
|
|||||||||||||
|
0.16–0.17 | 0.14 | 0.14–0.16 | 0.16 | |||||||||||||
|
0.14–0.18 | 0.14–0.16 | 0.13–0.17 | 0.14–0.17 | 0.16–0.17 |
|
|||||||||||
|
0.16–0.17 | 0.13–0.15 | 0.12–0.15 | 0.12–0.15 | 0.16–0.17 | 0.13–0.16 |
|
||||||||||
|
0.20 | 0.18 | 0.17–0.19 | 0.17–0.19 | 0.18 | 0.17–0.19 | 0.17–0.18 | ||||||||||
|
0.15–0.16 | 0.13 | 0.13–0.15 | 0.13–0.15 | 0.15 | 0.12–0.15 | 0.13–0.14 | 0.16 | |||||||||
|
0.18–0.19 | 0.16 | 0.16–0.17 | 0.16–0.18 | 0.18 | 0.17–0.18 | 0.15–0.16 | 0.17 | 0.15 | ||||||||
|
0.14–0.16 | 0.14 | 0.14–0.15 | 0.11–0.13 | 0.16 | 0.14–0.16 | 0.13–0.14 | 0.17 | 0.12 | 0.17 | |||||||
|
0.15–0.17 | 0.16 | 0.15–0.16 | 0.16–0.19 | 0.19 | 0.16–0.18 | 0.15–0.17 | 0.15 | 0.14 | 0.18 | 0.16 | ||||||
|
|
0.15–0.16 | 0.14–0.18 | 0.14–0.18 | 0.16–0.18 | 0.13–0.18 | 0.14–0.18 | 0.18–0.2 | 0.13–0.17 | 0.17–0.19 | 0.15–0.17 | 0.17–0.21 |
|
||||
|
0.20–0.21 | 0.18 | 0.18–0.19 | 0.20–0.21 | 0.18 | 0.18–0.20 | 0.18 | 0.20 | 0.18–0.19 | 0.19 | 0.19 | 0.20 | 0.19–0.22 | ||||
|
0.15 | 0.14 | 0.14–0.17 | 0.13–0.15 | 0.15 | 0.09–0.14 | 0.13–0.14 | 0.15 | 0.11 | 0.16 | 0.13 | 0.16 | 0.13–0.16 | 0.16 | |||
|
0.2–0.21 | 0.19 | 0.18–0.20 | 0.17–0.20 | 0.19 | 0.18–0.20 | 0.19 | 0.18–0.19 | 0.18 | 0.2–0.21 | 0.19 | 0.18 | 0.19–0.2 | 0.21 | 0.19 | ||
|
0.16–0.18 | 0.17 | 0.16–0.19 | 0.16–0.19 | 0.17–0.18 | 0.15–0.19 | 0.17–0.18 | 0.16–0.17 | 0.16 | 0.19–0.21 | 0.17 | 0.14–0.16 | 0.16–0.2 | 0.19–0.2 | 0.15–0.16 |
|
|
|
|||||||||||||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
||||||||||||||||
|
0.08–0.1 | ||||||||||||||||
|
0.08–0.1 | 0.06–0.07 |
|
||||||||||||||
|
0.06–0.11 | 0.06–0.08 | 0.03–0.06 |
|
|||||||||||||
|
0.09–0.11 | 0.08 | 0.06–0.07 | 0.07–0.08 | |||||||||||||
|
0.07–0.11 | 0.07–0.08 | 0.04–0.06 | 0.05–0.09 | 0.06–0.07 |
|
|||||||||||
|
0.07–0.08 | 0.06–0.07 | 0.04–0.05 | 0.04–0.06 | 0.06 | 0.06–0.08 |
|
||||||||||
|
0.08–0.11 | 0.07–0.1 | 0.06–0.08 | 0.05–0.08 | 0.07–0.09 | 0.06–0.09 | 0.06–0.08 |
|
|||||||||
|
0.07–0.11 | 0.05–0.07 | 0.03–0.06 | 0.04–0.07 | 0.04–0.08 | 0.05–0.07 | 0.04–0.07 | 0.04–0.09 |
|
||||||||
|
0.08–0.1 | 0.05 | 0.06–0.07 | 0.05–0.07 | 0.08 | 0.07 | 0.06 | 0.07–0.09 | 0.05–0.07 | ||||||||
|
0.07–0.08 | 0.07 | 0.05 | 0.05–0.07 | 0.08 | 0.06–0.07 | 0.05–0.06 | 0.05–0.07 | 0.05–0.07 | 0.07 | |||||||
|
0.06–0.07 | 0.06 | 0.04–0.05 | 0.03–0.06 | 0.06 | 0.05–0.06 | 0.04 | 0.05–0.07 | 0.04–0.06 | 0.06 | 0.05 | ||||||
|
|
0.07–0.08 | 0.06–0.07 | 0.05–0.08 | 0.07–0.08 | 0.05–0.07 | 0.06–0.07 | 0.06–0.08 | 0.06–0.08 | 0.07–0.08 | 0.05–0.06 | 0.05–0.06 |
|
||||
|
0.07–0.1 | 0.08 | 0.06–0.08 | 0.06–0.08 | 0.08–0.09 | 0.06–0.09 | 0.06–0.08 | 0.04–0.07 | 0.05–0.08 | 0.07–0.08 | 0.05–0.06 | 0.06–0.07 | 0.06–0.08 |
|
|||
|
0.09–0.1 | 0.09 | 0.08 | 0.07–0.1 | 0.08 | 0.07–0.08 | 0.08 | 0.08–0.1 | 0.06–0.08 | 0.1 | 0.08 | 0.07 | 0.07–0.08 | 0.08–0.09 | |||
|
0.1–0.11 | 0.09 | 0.07–0.09 | 0.07–0.09 | 0.08 | 0.08 | 0.08 | 0.08–0.09 | 0.06–0.08 | 0.08 | 0.07–0.08 | 0.08 | 0.08 | 0.07–0.09 | 0.08–0.09 | ||
|
0.09–0.11 | 0.07 | 0.08–0.09 | 0.07–0.09 | 0.09 | 0.07–0.09 | 0.08 | 0.07–0.08 | 0.06–0.09 | 0.08 | 0.08 | 0.08 | 0.08–0.09 | 0.06–0.08 | 0.09 |
|
BI phylogeny showing inferred phylogenetic relationships of old world racers. Labelled in pink color are the sequences generated in this study, colored blue is the sample from this study+GenBank sequence. Posterior probability support values shown at internal branches, values below 70 are not shown.
Maximum Likelihood phylogeny constructed using
Table with summarized characters used for osteological comparison of
|
|
|
|
|
|
|
|
|
|
individually short or long, one tip | short, in contact with nasals | 0.93–1.04 | 40%–50% | present only in 60% of examined specimens | individually unstructured or structured | 1.21–1.59 | 0.73–0.95 | 0.81–1.01 | |
|
short, one tip | short, in contact with nasals | 1.16–1.18 | 47%–58% | present | individually unstructured or structured | 1.54 3 | 0.853 | 1.023 |
|
long, two tips | short, in contact with nasals | 1.24 | 68% | absent | structured | 1.70 [1.32–1.48] | 0.85 [0.68–0.79] | 1.15 [0.95–1.05] |
|
long, one tip or stout | long, not in contact with nasals | 1.27–1.39 | 60%–77% | absent | structured | 1.60–1.70 [1.41–1.55] | 0.83–0.88 [0.55–0.64] | 1.20–1.27 [1.09–1.25] |
|
long, stout | long, in contact with nasals | 1.28 | 56% | absent | unstructured | 1.68 [1.49–1.51] | 0.86 [0.76–0.78] | 1.10 [1.16–1.18] |
|
/ | / | / | / | / | / | [1.22–1.39] | [0.63–0.71] | [0.84–1.07] |
|
/ | / | / | / | / | / | 1.61 [1.20–1.41] | 0.86 [0.66–0.70] | 0.80 [0.87–1.16] |
|
/ | / | / | / | / | / | 1.08 | 0.62 | 0.80 |
Segment plots of characters related to middorsal vertebrae values between
Ratio of the length of centrum to the width across prezygapophyses between outer edges of articular facets (LC/WP).
Ratio of the length of centrum and the least width of neural arch (LC/WN).
Ratio of length of neural spine to least width of neural arch (LN/WN).
Gazetteer of confirmed locality records for
Locality verbatim | Current locality | State | Country | Latitude | Longitude | elevation in m | References |
---|---|---|---|---|---|---|---|
Casemcottah [type locality] | Kasimkota | Andhra Pradesh | India | 17.66.660 | 82.967.483 | 26 | |
Ellore | Eluru | Andhra Pradesh | India |
|
|
17 | |
Nallamala Hills | Andhra Pradesh | India |
|
|
350 | ||
Pulicat Lake | Andhra Pradesh | India |
|
|
1 | ||
Rajamundry North Circars | Rajamundri | Andhra Pradesh | India |
|
|
67 | |
Rollapenta | Rollapenta | Andhra Pradesh | India |
|
|
474 | |
Sundipentha/Sikharam | Sundipenta | Andhra Pradesh | India |
|
|
470 | |
Sunnipenta, Nagarjunasagar Srisailam Tiger Reserve | Sunnipenta | Andhra Pradesh | India |
|
|
414 | |
Thummalapalle uranium mining station, Kadapa and Anantapur districts | Andhra Pradesh | India |
|
|
250 | ||
Vizagapatam | Visakhapatnam | Andhra Pradesh | India |
|
|
26 | |
Monghyr | Munger | Bihar | India |
|
|
54 | |
Pancht Hill, 2.5 miles from Inanpur, Manbhum | Bihar | India |
|
|
213 | ||
Patna | Patna | Bihar | India |
|
|
60 | |
Tinpahar, Rajmahal | Rajmahal | Bihar | India |
|
|
34 | |
Kaberdham | Kabirdham | Chhatisgarh | India |
|
|
365 | This study. |
Pharsabhar block | Jashpur | Chhatisgarh | India |
|
|
504 | |
Bondla Wildlife Sanctuary | Bondla Wildlife Sanctuary | Goa | India |
|
|
199 | |
Goa | Goa City | Goa | India |
|
|
90 | Bethencourt Ferreira (1897), |
Alcock Ashdown & Co., Old Port Bhavnagar | Gujarat | India |
|
|
40 | ||
Baroda | Vadodara | Gujarat | India |
|
|
41 | |
Bharuch | Bharuch | Gujarat | India |
|
|
22 | |
Bhavnagar | Bhavnagar | Gujarat | India |
|
|
32 | |
Dahod City | Dahod | Gujarat | India |
|
|
309 | |
Dangs | Dang | Gujarat | India |
|
|
352 | |
Gandhinagar | Gandhinagar | Gujarat | India |
|
|
82 | |
Gir forest | Gir forest | Gujarat | India |
|
|
278 | |
Girnar Hill | Gujarat | India |
|
|
409 |
|
|
Kala Gadba | Gujarat | India |
|
|
201 | ||
Mehsana | Mehsana | Gujarat | India |
|
|
86 | |
near Mahal | Mahal | Gujarat | India |
|
|
318 | |
Panchmahals | Panchmahal | Gujarat | India |
|
|
146 | |
Sabarkantha | Sabarkantha | Gujarat | India |
|
|
209 | |
Samot village, Shoolpaneshwar WS | Samot village, Shoolpaneshwar WS | Gujarat | India |
|
|
199 | |
Saurashtra | Gujarat | India |
|
|
122 | Patel & Vyas (2019). | |
Surat, suburban residential complex | Gujarat | India |
|
|
8 | ||
Valsad | Valsad | Gujarat | India |
|
|
28 | |
Vansda National Park | Gujarat | India |
|
|
144 | ||
Vizapur | Vijapur | Gujarat | India |
|
|
126 | |
Ambala | Ambala | Haryana | India |
|
|
281 | |
Hazaribag district | Jharkhand | India |
|
|
608 | Prakash & Raziuddin (2009). | |
Balekola, Mysore | Bealgola | Karnataka | India |
|
|
751 | Aengals & Pradhan (2013). |
Belgaum | Belgaum | Karnataka | India |
|
|
759 | |
Bengaluru | Bengaluru | Karnataka | India |
|
|
924 | This study. |
Chincholi Forest, Kalaburagi district | Karnataka | India |
|
|
617 | ||
Coorg | Coorg | Karnataka | India |
|
|
951 | |
Hosar, near Bangalore | Karnataka | India |
|
|
873 | ||
Mysore | Mysuru | Karnataka | India |
|
|
752 | |
Collagelly Hills, Coimbatore dist | Kollegal Hills | Karnataka | India |
|
|
644 | |
Shivamogga City Corporation | Karnataka | India |
|
|
575 | ||
Muthanga Wildlife Sanctuary | Muthanga, Wayanad district | Kerala | India |
|
|
868 | Radhakrishnan (1998) |
Barkatullah University, Bhopal | Madhya Pradesh | India |
|
|
481 | ||
Bhopal district | Madhya Pradesh | India |
|
|
497 | ||
Dewas | Dewas | Madhya Pradesh | India |
|
|
540 | |
Dumna Nature Park, Jabalpur | Madhya Pradesh | India |
|
|
462 | ||
Gwalior | Gwalior | Madhya Pradesh | India |
|
|
220 | |
Indore | Indore | Madhya Pradesh | India |
|
|
554 | |
Jabalpur | Jabalpur | Madhya Pradesh | India |
|
|
420 | |
Kanha National Park | Kanha National Park | Madhya Pradesh | India |
|
|
602 | Argawal (1976), |
Katra Hills, Bhopal | Madhya Pradesh | India |
|
|
472 | ||
Mandla | Mandla | Madhya Pradesh | India |
|
|
450 | |
Pachmarhi Biosphere Reserve | Madhya Pradesh | India |
|
|
1053 | ||
Satpura Tiger Reserve | Madhya Pradesh | India |
|
|
968 | ||
Saugor | Sagar | Madhya Pradesh | India |
|
|
519 | |
Shahdol | Shahdol | Madhya Pradesh | India |
|
|
510 | |
Sita Hill, Jabalpur | Sita Hill, Jabalpur | Madhya Pradesh | India |
|
|
423 | Sharma, R.C. (1982). |
Sonemuda (Anuppur District) | Madhya Pradesh | India |
|
|
972 | Ingle (2020). | |
Ujjain | Ujjain | Madhya Pradesh | India |
|
|
494 | |
Ahmednagar | Ahmednagar | Maharashtra | India |
|
|
855 | |
Akola district | Maharashtra | India |
|
|
289 | ||
Ale village, ca. 20 km N of Narayangon | Ale, Pune | Maharashtra | India |
|
|
668 | Sharma, R. C. (1982). |
Atpadi Tahasil, Sangli district | Maharashtra | India |
|
|
573 | ||
Aurangabad | Aurangabad | Maharashtra | India |
|
|
111 | |
Bassein Fort, Thane | Maharashtra | India |
|
|
10 | ||
Bhanadara, Nagpur | Bhanadara, Nagpur | Maharashtra | India |
|
|
263 | This study. |
Bohali | Bohali | Maharashtra | India |
|
|
495 | |
Bombay Presidency | Mumbai | Maharashtra | India | ||||
Borivali (Gorai) Mangroves | Maharashtra | India |
|
|
91 | ||
Buldhana district, Amravati division | Maharashtra | India |
|
|
473 | ||
Chink Hill | Chink Hill | Maharashtra | India |
|
|
459 | |
Dahanu Forest Division | Maharashtra | India |
|
|
15 | ||
Deolali | Deolali, Nasik | Maharashtra | India |
|
|
555 | |
Devagad, Sindhudurg district | Maharashtra | India |
|
|
50 | ||
Ghorawadi, near Talegaon, Pune district | Talegaon, Pune | Maharashtra | India |
|
|
599 | |
Jalgaon district | Maharashtra | India |
|
|
218 | ||
Juvem, Salsette | Salsette, Mumbai | Maharashtra | India |
|
|
171 | |
Kaas Plateau | Kas plateau, Satara district | Maharashtra | India |
|
|
1191 | |
Katraj Snake Park Pune | Maharashtra | India |
|
|
665 | ||
Khandalla | Khandala | Maharashtra | India |
|
|
297 | |
Kolhapur district | Maharashtra | India |
|
|
570 | ||
Kurduvadi | Kurduvadi | Maharashtra | India |
|
|
516 | |
Maharashtra Nature Park | Mumbai | Maharashtra | India |
|
|
6 | |
Malegaon Tehsil, Washim district | Maharashtra | India |
|
|
432 | ||
Mokhada and Jawhar, Palghar district | Maharashtra | India |
|
|
353 | ||
Nagar | Nagarm, Ahmednagar | Maharashtra | India |
|
|
734 | |
Nagpur | Nagpur | Maharashtra | India |
|
|
344 | |
Nanded city | Maharashtra | India |
|
|
388 | ||
Navegaon NP, Gondia district | Maharashtra | India |
|
|
331 | ||
Palghar region, Thane | Maharashtra | India | 19.6978751 | 19.6978751 | 47 | ||
Panchgani | Panchgani, Satara | Maharashtra | India |
|
|
1258 | |
Parel, Bombay | Parel, Mumbai | Maharashtra | India |
|
|
5 | |
Parvati-Pachgaon Hills, Pune | Pune | Maharashtra | India |
|
|
606 | |
Poona | Pune | Maharashtra | India |
|
|
556 | |
Powai, Salsette | Powai, Mumbai | Maharashtra | India |
|
|
43 | |
Puna | Pune | Maharashtra | India |
|
|
556 | |
Ranidoh region, Pench NP | Pench NP | Maharashtra | India |
|
|
551 | |
Santa Cruz, Salsette | Salsette, Mumbai | Maharashtra | India |
|
|
171 | |
Swami Ramanand Teerth Marathwada University, Nanded | Maharashtra | India |
|
|
384 | ||
Thana | Thane | Maharashtra | India |
|
|
20 | |
Uran wetland, Navi, Mumbai | Mumbai | Maharashtra | India |
|
|
8 | |
Vidarbha region | Maharashtra | India |
|
|
291 | ||
Vidyanagari campus Mumbai Uni, Maharashtra | Vidyanagari Campus | Maharashtra | India |
|
|
258 | |
Visapur | Visapur, Ahmednagar | Maharashtra | India |
|
|
115 | |
Yavatmal | Yavatmal | Maharashtra | India |
|
|
377 | |
Marol, Mumbai | Mumbai | Mahrashtra | India |
|
|
15 | |
Baripada | Baripada | Odisha | India |
|
|
64 | This study. |
Choudwar, Cuttack | Choudwar, Cuttack | Odisha | India |
|
|
32 | This study. |
Gandhamardan Hills | Odisha | India |
|
|
293 | ||
Nandankanan Biological Park | Nandankanan Biological Park. Cuttack | Odisha | India |
|
|
36 | |
Patharakali area, Bhadrak district | Odisha | India |
|
|
19 | ||
Pondicherry | Pondicherry | Puducherry | India |
|
|
14 | This study. |
near Hemavas Dam, Pali district | Hemwas, Pali | Rajasthan | India |
|
|
220 | |
Pratapgarh district | Rajasthan | India |
|
|
459 | Anonymous (n. d., p. 13) | |
Ramri | Rajasthan | India |
|
|
253 | ||
Farahabad | Telangana state | India |
|
|
537 |
|
|
Hyderabad Region | Telangana state | India |
|
|
486 | Walmiki) 2012b), |
|
Ippalapally | Telangana state | India |
|
|
419 | ||
Jannaram | Telangana state | India |
|
|
190 | ||
Kerimeri | Telangana state | India |
|
|
277 | ||
Khanapur | Telangana state | India |
|
|
248 | ||
Kinnerasani | Telangana state | India |
|
|
94 | ||
Old Bowenpally | Telangana state | India |
|
|
548 | ||
OU Tarnaka | Telangana state | India |
|
|
517 | ||
Rushulacheruvu | Telangana state | India |
|
|
354 | ||
Vijayapuri | Telangana state | India |
|
|
149 | ||
Allahabad | Prayagraj | Uttar Pradesh | India |
|
|
103 | |
Benares | Varanasi | Uttar Pradesh | India |
|
|
79 | |
Fyzabad | Faizabad | Uttar Pradesh | India |
|
|
105 | |
Mathura region | Uttar Pradesh | India |
|
|
165 | ||
Calcutta | Kolkata | West Bengal | India |
|
|
10 | |
Dum Dum | West Bengal | India | 22.647.051 | 88.431.686 | 10 | ||
Durgapur city | Durgapur | West Bengal | India | 23,5788078 | 87,3264641 | 88 | |
Hosur Hills | Tamil Nadu | India |
|
|
851 | ||
Tambaram, Chennai | Tamil Nadu | India |
|
|
36 | ||
Tambaram, Chennai | Tamil Nadu | India |
|
|
36 | ||
Lal Suhanra National Park, Bahawalpur | Bahawalpur | Punjab | Pakistan |
|
|
125 | |
Badin | Badin | Sindh | Pakistan |
|
|
10 | |
Hala | Hala | Sindh | Pakistan |
|
|
25 | |
Jati | Jati | Sindh | Pakistan |
|
|
-50 | |
Jogis | Jogi | Sindh | Pakistan |
|
|
50 | |
Larkana district | Larkana | Sindh | Pakistan |
|
|
48 | |
Makli Hills near Tatta | Makli hills nr. Tatta | Sindh | Pakistan |
|
|
30 | |
Mohenjo-daro | Mohenja-daro | Sindh | Pakistan |
|
|
52 | |
Pir Patho | Pir Patho, Thatta | Sindh | Pakistan |
|
|
20 | |
Raj Malk, Thatta district, Mirpur Sakro Area | Raj Malk, Thatta | Sindh | Pakistan |
|
|
7 | |
Sonda (1 mile east of) | Sonda, Thatta district | Sindh | Pakistan |
|
|
15 | |
Tatta (4 miles east of) | Thatta | Sindh | Pakistan |
|
|
10 | |
|
|||||||
Anaikatty, Coimbatore | Anaikatti, Coimbatore | Tamil Nadu | India |
|
|
582 | This study. |
Anamalai Hills | Anamalai hills | Tamil Nadu | India |
|
|
404 | |
Auroville, Villupuram | Auroville, Villupuram | Tamil Nadu | India |
|
|
52 | |
Batlagundu, Dindugal district | Batlagundu, Dindugal | Tamil Nadu | India |
|
|
237 | This study. |
Coimbatore | Coimbatore `by implication’ | Tamil Nadu | India |
|
|
415 | |
Coutrallam | Courtallam | Tamil Nadu | India |
|
|
190 | This study. |
Karur | Karur | Tamil Nadu | India |
|
|
133 | This study. |
Madurai | Madurai | Tamil Nadu | India |
|
|
139 | |
Manimutharu | Tirunelveli | Tamil Nadu | India |
|
|
79 | This study. |
Maruthuvazhmalai, Kanyakumari | Maruthuvazhmalai, Kanyakumari | Tamil Nadu | India |
|
|
243 | |
Meghamalai Hills | Meghamalai Hills | Tamil Nadu | India |
|
|
445 | |
Pollachi I | Pollachi | Tamil Nadu | India |
|
|
293 | This study. |
Pollachi II | Pollachi | Tamil Nadu | India |
|
|
342 | This study. |
Salem | Salem | Tamil Nadu | India |
|
|
314 | This study. |
Tiruneveli | Tirunelveli | Tamil Nadu | India |
|
|
44 | |
Trichinopoly | Tiruchirapalli | Tamil Nadu | India |
|
|
84 | |
Tuticorin (type locality) | Thoothukudi | Tamil Nadu | India |
|
|
7 | |
Vadipatti, Madurai dist | Vadipatti, Madurai dist | Tamil Nadu | India |
|
|
214 | This study. |
(Material examined for this study. Symbol * denotes for specimens CT scanned for analysis of the osteology and symbol ° for specimens used in analysis of dorsal scale reduction.)
Without specified locality:
INDIA – without specified locality, Russell’s dried snake skin collection:
INDIA – Andhra Pradesh:
EGYPT –
INDIA –Tamil Nadu:
PAKISTAN – Sindh:
INDIA – Andhra Pradesh:
TURKMENISTAN –
PAKISTAN – Sindh:
Without specified locality: