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Research Article
First description of the female of Achalinus sheni (Serpentes, Xenodermidae), with expanded description of this species
expand article infoYi-han Ma‡, Jia-yu Liu‡, Hui Li‡, Lin Zhou§, Yu-hao Xu|, De-yong Peng, Zhi-qiang Zhang§, Xiao-yang Mo‡#
‡ Hunan Normal University, Changsha, China
§ Central South University of Forestry and Technology, Changsha, China
| Qinghai University, Xining, China
¶ Jintong Mountain Nature Reserve Management Office, Hunan Nanshan National Park Administration, Shaoyang, China
# Huaihua University, Huaihua, China

Corresponding author: Zhi-qiang Zhang ( csfuzzq@126.com )

Corresponding author: Xiao-yang Mo ( moxiaoyang@hunnu.edu.cn )

Academic editor: Silke Schweiger
© 2024 Yi-han Ma, Jia-yu Liu, Hui Li, Lin Zhou, Yu-hao Xu, De-yong Peng, Zhi-qiang Zhang, Xiao-yang Mo.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Ma Y-h, Liu J-y, Li H, Zhou L, Xu Y-h, Peng D-y, Zhang Z-q, Mo X-y (2024) First description of the female of Achalinus sheni (Serpentes, Xenodermidae), with expanded description of this species. Herpetozoa 37: 189-195. https://doi.org/10.3897/herpetozoa.37.e124331
ZooBank: urn:lsid:zoobank.org:pub:01520209-7D04-4286-9697-9141EF343B1D
Open Access

Abstract

Achalinus sheni Ma, Xu, Qi, Wang, Tang, Huang & Jiang, 2023 was originally described based on only male specimens from Lianyuan City and Nanyue District, Hunan, China. So far, no information on the females of this species is available. After molecular systematics and morphological characters of recently collected specimens from Xinshao County, Hunan, several specimens (3 males and 1 female) were identified as this species. Therefore, we provided supplementary descriptions of the female characteristics of this species in this study. Meanwhile, we extended the description of this species.

Key Words

Morphology, Phylogenetics, Xinshao County

Introduction

Achalinus Peters, 1869, commonly known as the odd-scaled snakes, is the most diverse genus of the family Xenodermidae (Uetz et al. 2022). It contains 28 recognized species, and is widely distributed in eastern and southeastern Asia, ranging from northern Vietnam to southwestern China, and partly into Japan, from which 21 species occur in China. In recent years, with extensive sampling and molecular phylogenetic methods, more than 20 members of the genus Achalinus have been discovered and described (Wang et al. 2019; Ziegler et al. 2019; Li et al. 2020; Luu et al. 2020; Miller et al. 2020; Hou et al. 2021; Huang et al. 2021; Li et al. 2021; Ha et al. 2022; Yang et al. 2022; Ma et al. 2023a, 2023b; Yang et al. 2023; Zhang et al. 2023; Li et al. 2024).

Achalinus sheni was described based on five specimens collected in Hunan Province, China: four male specimens (ANU20230012—ANU20230015) from Lianyuan City and one male specimen (CIB 119043) from Nanyue District (Ma et al. 2023b). Molecular phylogeny inferred from the mitochondrial COⅠ gene fragment revealed that this new species is most closely related to A. yunkaiensis, but it can be distinguished from A. yunkaiensis by the following morphological characters: (1) relative length of supraocular and upper anterior temporal (supraocular equal to or longer than anterior temporal, SPOL/ATUL 0.99–1.20 vs. supraocular shorter than anterior temporal, SPOL/ATUL 0.55–0.83); (2) higher ventral + subcaudals counts in males (220–225 vs. 200–212); (3) higher ventral scale counts in males (161–170 vs. 150–162); (4) higher subcaudal scale counts in males (55–61 vs. 49–56). However, to date there have been no morphological comparisons between females of these two species.

During a herpetofaunal survey of Hunan, China in 2022, one female and three male Achalinus were collected from Xinshao County, Shaoyang City (Fig. 1). Morphological and molecular phylogenetic analyses revealed that the specimens are A. sheni. Here we provide morphological data for the newly collected female specimen of A. sheni , and extend the description of this species.

Figure 1. 

Sampling sites of Achalinus sheni: solid red star, the locality of the holotype; solid red circle, the locality of the paratype; solid green triangle, the locality of the newly collected female specimen.

Materials and methods

Sampling

Four odd‑scaled snake specimens were collected from Xinshao County, Shaoyang City, Hunan Province, China (HNNU2022001—HNNU2022004). The four specimens were fixed in 75% ethanol and deposited in the Vertebrate Zoology Laboratory, College of Life Science, Hunan Normal University.

Morphological examination

Morphological descriptions followed Zhao (2006) and Ma et al. (2023b). Abbreviations in this study are as follows: snout-vent length (SVL): snout-vent length from tip of snout to anterior margin of the cloaca; tail length (TaL): tail length from posterior margin of cloaca to tip of tail; total length (TL): from snout tip to tail end; head length (HL): from the tip of snout to the posterior margin of mandible; head width (HW): from the widest part of the head in dorsal view; eye horizontal diameter (ED): from the most anterior edge of the eye to the most posterior edge; loreal height (LorH): measured from the highest point to the lowest point of the loreal in lateral view; loreal length (LorL): from the most anterior edge of the loreal to the most posterior edge of the loreal in lateral view; length of the suture separating the internasals (LSBI); length of the suture separating the prefrontals (LSBP); length of the supraocular (SPOL): horizontal distance between the anterior and posterior tips of the supraocular, and length of the upper anterior temporal (ATUL): horizontal distance between the anterior and posterior tips of the upper anterior temporal. We also directly compared the length of the sutures between the internasals and prefrontals (LSBI vs. LSBP).

The morphological features and their abbreviations are as follows: loreals (Loreal), supralabials (SPL), infralabials (IFL), the number of infralabials touching the first pair of chin shields (IFL-1st Chin), supraoculars (SPO), temporals (TEM), the number of anterior temporals touching the eye (aTEM-Eye), ventral scales (VEN), subcaudals (SC), cloacal plate entire or divided (CP), dorsal scale rows (DSR) (counted at one-head-length posterior of head, at midbody, at one-head-length anterior of cloacal plate). Bilateral scale counts were given as left/right.

Phylogenetic analyses

Genomic DNA was extracted from preserved liver tissue using the TIANamp Genomic DNA Kit. The fragment of the mitochondrial DNA gene encoding cytochrome c oxidase subunit Ⅰ (COⅠ) was amplified using the primer pairs Chfm4 and Chrm4 (Che et al. 2012). The PCR products were sequenced at Shanghai Map Biotech Co., Ltd. The homologous sequences of the Achalinus species and the outgroups species were downloaded from GenBank (Table 1).

Table 1.
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Localities, voucher information and GenBank numbers for all samples used in this study.

Species Locality Voucher NO. Accession
A. sheni Lianyuan, Hunan, China ANU20230013 OR178146
A. sheni Lianyuan, Hunan, China ANU20230014 OR178147
A. sheni Nanyue, Hunan, China CIB 119043 OR189183
A. sheni Xinshao, Hunan, China HNNU2022001 PP854453
A. ater Huaping Nature Reserve, Guangxi, China SYSr00852 MN380334
A. dabieshanensis Fuziling Provincial Reserve, Anhui, China AHU2018EE0710 MW316598
A. damingensis Shanglin, Nanning, Guangxi, China ANU20220009 OP644487
A. dehuaensis Dehua, Fujian, China YBU13013 MZ442642
A. emilyae HoanhBo, Quang Ninh, Vietnam IEBR4465 MK330857
A. formosanus Taiwan, China RN2002 KU529452
A. huangjietangi Huangshan, Anhui, China HSR18030 MT380191
A. hunanensis Huaihua, Hunan, China CIB119039 OQ848425
A. juliani HaLang, Cao Bang, Vietnam IEBRA.2018.8 MK330854
A. meiguensis Mianyang, Sichuan, China GP835 MZ442641
A. nanshanensis Nanshan National Park, Hunan, China HNNU230903 OR523370
A. niger Taiwan, China RN0667 KU529433
A. ningshanensis Ningshan, Shaanxi, China ANU20220006 ON548422
A. panzhihuaensis Yanbian, Sichuan, China KIZ040189 MW664862
A. pingbianensis Honghe, Yunnan, China YBU18273 MT365521
A. quangi northern Vietnam sp4 OQ197471
A. rufescens Hongkong, China SYSr001866 MN380339
A. spinalis Badagong Mountains, Hunan, China SYSr001327 MN380340
A. timi ThuanChau, Son La, Vietnam IEBRA.2018.10 MK330856
A. tranganensis NinhBinh, Vietnam VNUFR.2018.21 MW023086
A. vanhoensis VanHo, Son La, Vietnam VNUFR.2019.13 ON677935
A. yangdatongi Wenshan Nature Reserve, Yunnan, China KIZ034327 MW664865
A. yunkaiensis Dawuling Forestry Station, Guangdong, China SYSr001443 MN380329
A. yunkaiensis Dawuling Forestry Station, Guangdong, China SYS r001502 MN380330
A. yunkaiensis Dawuling Forestry Station, Guangdong, China SYS r001503 MN380331
A. zugorum Bac Me, Ha Giang, Vietnam IEBR4698 MT502775
Fimbrios klossi Quang Ngai, Vietnam IEBR3275 KP410744
Parafmbrios lao Louangphabang, Laos MNHN2013.1002 KP410746
Xenodermus javanicus Sumatera Barat, Indonesia — KP410747

The COⅠ sequences (639 bp) were assembled using SeqMan in the DNASTAR software package (Burland, 2000), and compared and aligned using MEGA 7 software (Kumar et al. 2018). The uncorrected pairwise distances (p-distance) were calculated in MEGA 7. Maximum likelihood analysis (Nguyen et al. 2015) was executed using IQ‑TREE 2 under the best-fit model TIM3 + F + I + G4 selected by Modelfinder according to AIC. Nodal support was estimated by 1,000 bootstrap replicates using the ultrafast bootstrap feature.

Results

Maximum likelihood trees showed consistent topology. The newly collected female specimen clustered with the specimens (including the holotype) of Achalinus sheni and demonstrated strong support (ML = 97, Fig. 2). The genetic distance (uncorrected p-distance) between the newly collected female specimen and the specimens (including the holotype) of A. sheni was only 0.7–0.8% (Table 2).

Figure 2. 

Phylogenetic tree of the genus Achalinus inferred from COⅠ gene fragments (629 bp) using Maximum Likelihood. The numbers above the branches represent the supporting values.

Table 2.
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Uncorrected p-distances (%) among Achalinus species inferred from mitochndrial COⅠ gene.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
1 HNNU2022001
2 A. sheni 0.7
3 A. sheni 0.7 0.0
4 A. sheni 0.8 0.2 0.2
5 A. ater 13.5 13.2 13.2 13.2
6 A. dabieshanensis 15.3 15.8 15.8 15.6 14.7
7 A. damingensis 14.4 14.0 14.0 14.0 8.0 15.8
8 A. dehuaensis 13.9 13.9 13.9 13.8 17.1 18.4 16.1
9 A. emilyae 14.2 13.7 13.7 13.5 11.9 17.7 13.4 15.7
10 A. formosanus 13.2 13.0 13.0 13.0 14.4 19.0 14.9 16.4 14.0
11 A. huangjietangi 13.6 13.6 13.6 13.3 15.0 8.9 16.3 16.5 14.5 15.6
12 A. hunanensis 12.5 12.1 12.1 12.3 7.3 16.9 6.1 14.9 13.2 13.7 16.8
13 A. juliani 14.4 14.0 14.0 14.2 7.0 15.8 8.7 15.1 12.9 12.5 14.6 8.8
14 A. meiguensis 14.6 14.1 14.1 13.9 15.4 17.7 16.8 18.1 15.4 15.6 15.2 16.4 16.8
15 A. nanshanensis 13.5 13.9 13.9 14.0 6.7 16.0 5.4 14.7 13.4 15.1 16.6 4.7 8.2 17.7
16 A. niger 12.7 12.5 12.5 12.5 13.5 15.8 14.4 16.2 12.4 9.2 13.9 13.2 12.5 13.9 13.5
17 A. ningshanensis 14.5 14.1 14.1 14.2 7.6 17.2 7.8 16.5 14.1 14.8 17.2 3.4 9.6 17.0 5.1 14.6
18 A. panzhihuaensis 14.6 14.6 14.6 14.6 16.2 16.6 15.5 15.3 16.6 16.0 15.2 16.2 15.5 11.6 15.5 14.4 17.4
19 A. pingbianensis 11.5 11.5 11.5 11.6 11.6 15.3 10.9 15.0 12.8 14.5 13.0 11.1 1.0 16.8 11.5 11.8 11.7 14.9
20 A. quangi 15.1 14.5 14.5 14.3 12.2 18.1 13.2 15.9 3.3 14.2 15.0 13.2 12.9 15.2 13.0 11.9 13.4 16.9 13.8
21 A. rufescens 13.7 13.5 13.5 13.3 12.9 16.9 13.9 14.4 8.0 14.0 14.3 12.1 12.5 17.3 12.2 12.7 12.3 16.0 12.8 7.9
22 A. spinalis 12.0 11.5 11.5 11.3 15.4 16.6 15.4 14.4 14.4 14.0 13.4 13.9 14.4 16.0 14.4 13.7 15.6 15.8 13.2 14.0 13.0
23 A. timi 14.2 13.9 13.9 13.8 13.2 16.4 13.5 15.9 13.4 13.5 14.8 12.0 14.4 15.8 13.9 11.9 13.6 15.5 11.8 13.5 14.2 14.4
24 A. tranganensis 14.0 13.7 13.7 13.7 12.9 15.3 14.4 14.0 12.0 17.1 13.4 14.0 13.9 16.4 13.7 14.5 15.2 16.4 13.2 12.4 11.7 15.1 14.0
25 A. vanhoensis 14.1 13.8 13.8 13.6 13.1 15.5 12.6 16.0 12.2 14.0 14.6 11.5 13.6 15.6 12.4 12.8 12.1 15.5 10.8 12.4 13.8 12.9 5.2 13.3
26 A. yangdatongi 14.0 13.7 13.7 13.8 6.2 16.6 5.6 14.0 12.8 14.4 14.6 5.1 7.3 17.1 4.4 13.7 5.9 15.5 11.3 12.6 11.5 14.2 13.1 12.8 11.3
27 A. yunkaiensis 6.5 6.5 6.5 6.2 13.2 14.9 12.7 15.1 13.5 12.4 12.5 12.0 12.9 15.8 12.7 12.4 13.7 15.7 11.6 14.0 13.7 12.2 14.2 13.9 13.6 12.0
28 A. zugorum 10.9 10.9 10.9 10.8 13.7 15.3 12.9 14.7 12.5 13.4 4.3 11.8 13.4 15.0 13.0 13.2 12.8 15.3 10.8 13.2 13.7 13.4 13.5 12.0 11.9 12.2 11.2

Achalinus sheni Ma, Xu, Qi, Wang, Tang, Huang & Jiang, 2023

Specimen examined

HNNU2022001, adult female, collected by Hui Li, Lin Zhou on April 11, 2022, from Xinshao County (27°23'58"N, 111°33'44"E, 420 m a. s. l.), Shaoyang City, Hunan Province, China.

Description of the female specimen

Adult female with a total length of 345.1 mm (SVL 294.8 mm and TaL 50.3 mm), tail relatively short, TL/ToL 0.149, body slender and cylindrical. Head distinct from neck, rostral small, triangular, only the upper tip is visible from above. Head length 9.16 mm, head width 3.95 mm, HL/HW 2.32. Eyes small, eye width 1.10 mm. Length of the suture between the internasals (LSBI 1.40 mm) subequal to the length of the suture between the prefrontals (LSBP 1.31 mm). Frontal pentagonal pointed backwards, much shorter than the parietals; each parietal bordered with an elongated nuchal, with no preoculars or postoculars. Nostril in anterior portion of nasal scale, posterior margin of nostril with a distinct nostril cleft. A single loreal scale present, extending from the nasal to the eye, distinctly wider than high. Temporals 2+2+3. aTMPs elongated, upper aTMP much smaller than the lower aTMP; upper aTMP and lower aTMP in contact with eye, lower aTMP also in contact with parietal scale. Supralabials 6, 4th–5th contact the eye, the last much elongated. A single mental scale present. Two chin shields, the anterior pair longer than the posterior pair. Infralabials 5, the first contacting each other posterior to the mental and anterior to the 1st chin shields, 1st–3rd contact the 1st chin shields. Dorsal scales 23-23-23, strongly keeled, dorsum with an inconspicuous longitudinal vertebral stripe. Ventrals (VEN) 173. Subcaudals (SC) 45, not paired. Cloacal plate (CP) entire (Table 3).

Table 3.
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Main morphological characters of Achalinus sheni.

Voucher number HNNU2022001 HNNU2022002 HNNU2022003 HNNU2022004
Sex Adult female Adult male Adult male Adult male
SVL 294.8 270.8 239.9 211.5
TaL 50.34 74.3 67.7 59.2
TL 345.1 345.1 307.6 270.7
TaL/TL 0.149 0.215 0.220 0.219
HL 9.16 8.76 7.86 7.65
HW 3.95 4.01 3.94 3.26
ED 1.10 0.97 1.02 0.76
SPL 6 6 6 6
SPL-Eye 4th–5th 4th–5th 4th–5th 4th–5th
IFL 5 5 5/6 5
Chin 2 2 2 2
IFL-1stChin 1st–3rd 1st–3rd 1st–3rd 1st–3rd
Loreal 1 1 1 1
LorH 0.88 0.87 0.93 0.73
LorL 1.38 1.14 1.31 1.00
LorH / LorL 0.63 0.76 0.71 0.73
LSBI vs LSBP =1 =1 =1 =1
SPO 1 1 1 1
SPOL 1.49 1.15 1.15 1.33
TEM 2+2+3 2+2+3 2+2+3 2+2+3
aTEM-Eye 2 2 2 2
ATUL 1.63 1.41 1.71 1.46
SPOL / ATUL 0.91 0.82 0.67 0.91
DSR 23-23-23 23-23-23 23-23-23 23-23-23
VEN 173 156 156 158
SC 45 61 61 62
VEN+SC 218 217 217 220
CP Entire Entire Entire Entire

Coloration of the female specimen in life

Scales possess a subtle iridescent quality which gives the dorsum a distinctive reflective brownish-black appearance. Dorsum dark brown and the five innermost dorsal scale rows a little darker, forming an inconspicuous longitudinal vertebral line. Chin shields are tan. On the ventral surface, an off-white shade prevails, with the edges of the ventral scales gradually transitioning from gray-white to black. Ventral side of tail brownish.

Coloration in preservative

(Fig. 3) The dorsal surface of the body uniformly brownish-black, slightly tinged with iridescence and the longitudinal vertebral line appears a little darker. Chin shields light brown. Ventrals generally pale brown, darker on both sides, free margins of ventral scales grayish-white. Ventral surface of tail light brown.

Figure 3. 

Adult female (HNNU2022001) of Achalinus sheni. A. Dorsolateral view; B. Ventral view; C. The tail of the male and female; D. Dorsal head view; E. Ventral head view; F. Light side of head view; G. Ventral head view. Photos by Le-Qiang Zhu.

Extended diagnosis

Upon examining the additional adult males of Achalinus sheni from Xinshao County, it was observed that they possessed fewer ventral scales compared to specimens from the type locality. However, the number of subcaudal scales remained approximately the same for individuals from both locations. Upon inspection of adult female A. sheni collected in Xinshao County, it was noted that it exhibited an extremely short tail and consequently fewer subcaudals and more ventral compared to adult male individuals.

Comparison

Compared to male individuals of A. sheni and A. yunkaiensis, female individuals of both species exhibit similar differences in the number of ventral scales and subcaudals. Compared to males, both female A. yunkaiensis and female A. sheni have fewer sublabial scales, which is consistent with the common characteristics of this genus. However, the number of sublabial scales of A. sheni is even less than that of female A. yunkaiensis (Table 4).

Table 4
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Comparing the differences between Achalinus sheni and Achalinus yunkaiensis in different regions from a taxonomical perspective.

Species Achalinus sheni Achalinus yunkaiensis
Lianyuan City & Nanyue District Xinshao County Guangdong
Sex Males(n=5) Males (n=3) Female Female
SVL 121.8–292.2 211.5–270.8 294.8 204–386.3
TaL 27.2–80.3 59.2–74.3 50.3 52–72.8
TL 149.0–371.3 270.7–345.1 345.1 256–448.1
TaL/TL 0.183 ~ 0.224 0.215–0.220 0.149 0.156–0.204
HL 10.07–10.95 7.65–8.76 9.16 -
HW 5.96–7.25 3.26–4.01 3.95 -
ED 1.09–1.11 0.76–1.02 1.10 -
SPL 6 6 6 6
SPL-Eye 4th–5th 4th–5th 4th–5th 4th–5th
IFL 5 (rarely 6) 5 (rarely 6) 5 6
Chin 2 2 2 2
IFL-1stChin 1st–3rd 1st–3rd 1st–3rd 1st–3rd/4th
Loreal 1 1 1 1
LorH 0.69–0.93 0.73–0.93 0.88 0.74–1.2
LorL 1.29–1.71 1.00–1.31 1.38 1.51–2.2
LorH / LorL 0.53–0.57 0.71–0.76 0.63 0.49–0.55
LSBI vs LSBP =1 =1 =1 =1
SPO 1 1 1 1
SPOL 1.21–1.59 1.15–1.33 1.49 1.26–1.60
TEM 2+2+3 2+2+3 2+2+3 2+2+3/4
aTEM-Eye 2 2 2 2
ATUL 1.20–1.48 1.41–1.71 1.63 1.93–2.90
SPOL / ATUL 0.99–1.16 0.67–0.91 0.91 0.55–0.65
DSR 23-23-23 23-23-23 23-23-23 23-23-23
VEN 161–170 156–158 173 144–156
SC 55–61 61–62 45 51–55
VEN+SC 220–225 217–220 218 195–205
CP Entire Entire Entire Entire

Discussion

Only five male specimens of the Achalinus sheni have been described in previous studies. This study reports the first discovery of a female of the same species and provides a detailed description and photograph of the female. In terms of morphological characteristics, the male specimens collected in Xinshao County are close to the holotype specimen from Lianyuan City. Some differences were noted in the female specimen: Tail relatively short, TL/ToL 0.149, which was different from that of the male specimens (0.183–0.224); the tail of the female specimen is suddenly tapered, distinct from the elongated tail of the male specimens.

Xinshao County and Lianyuan City are geographically adjacent, and the climate is not obviously different. Achalinus sheni is fossorial and difficult to find. Only one female specimen was collected in this study. Whether the apparent differences between the sexes of this species are consistent, whether there are further differences, and the reasons for these differences have not yet been fully investigated and analyzed.

Author contributions

Yi-han Ma: Conceptualization, Writing - original draft. Jia-yu Liu: Data curation, Investigation. Hui Li: Data curation, Formal analysis. Lin Zhou: Investigation. Yu-hao Xu: Methodology. De-yong Peng: Investigation. Zhi-qiang Zhang: Funding acquisition, Methodology. Xiao-yang Mo: Writing - review and editing, Methodology.

Acknowledgements

We thank Le-qiang Zhu for taking photos of the specimen, and thank the driver Xian Zhang for his help in the field investigation process (Lutou Experimental Forest Farm, Central South University of Forestry and Technology).

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1 Yi-han Ma and Jia-yu Liu contributed equally as the first authors.
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