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Communication

First European Occurrence and Genetic Diversity of the Steppe Ribbon Racer Psammophis lineolatus (Brandt, 1838) (Serpentes: Psammophiidae)

by
Kazhmurat Akhmedenov
1,
Andrey Bakiev
1,2,
Anastasia Klenina
1,2,
Svetlana Lukonina
3 and
Evgeniy Simonov
4,*
1
Faculty of Natural Geography, M. Utemisov West Kazakhstan University, N. Nazarbayev Avenue 162, Uralsk 090000, Kazakhstan
2
Institute of Ecology of Volga River Basin, Samara Federal Research Center of Russian Academy of Sciences, Komzin Str. 10, Togliatti 445003, Russia
3
Department of Zoology and Ecology, Penza State University, Krasnaya Str. 40, Penza 440026, Russia
4
Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, Leninsky Avenue 33, Moscow 119071, Russia
*
Author to whom correspondence should be addressed.
Taxonomy 2025, 5(4), 54; https://doi.org/10.3390/taxonomy5040054
Submission received: 14 August 2025 / Revised: 2 October 2025 / Accepted: 8 October 2025 / Published: 10 October 2025
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Abstract

The recent update to the list of European herpetofauna species, published in 2020, includes 206 non-avian reptile species. The steppe ribbon racer, Psammophis lineolatus, is a snake species widely distributed across the deserts and semi-deserts of Central Asia and neighboring regions. Previously unrecorded in Europe, two specimens were discovered west of the Ural River during surveys in western Kazakhstan’s northeastern Caspian region. This is the first record of the species in Europe and marks its most northwestern distribution. DNA barcoding analysis revealed substantial mitochondrial diversity in the region, with European specimens having distinct haplotypes (0.73% p-distance), indicating colonization by two different phylogenetic lineages. This discovery highlights the need for further research on the P. lineolatus intraspecific diversity, phylogeography, and taxonomy.

1. Introduction

The latest update of the European herpetofauna species list, compiled by the Taxonomic Committee of the Societas Europaea Herpetologica, encompasses 95 amphibian species and 206 non-avian reptile species [1]. These numbers depend on the geographical boundaries that define Europe. For its classification, the committee references the 1997 and 2014 Atlases of Amphibians and Reptiles in Europe [2,3]. The Europe-Asia border, particularly around the Ural River and Ural Mountains, has historically been a point of contention. In 2020, the committee classified all territories west of these landmarks as European, including Russia’s Yekaterinburg Region. This is consistent with the widespread recognition of the Ural River as the Europe-Asia boundary by various authorities, e.g., [4].
One of the Asian species known to approach but not cross the Ural River boundary is the steppe ribbon racer Psammophis lineolatus (Brandt, 1838). This species is widely distributed across the deserts and semi-deserts of the Turanian region and neighboring areas. It is found along the east shore of the Caspian Sea, through Kazakhstan, Turkmenistan, Iran, Uzbekistan, Afghanistan, Kyrgyzstan, Tajikistan, and Pakistan, and as far as southwestern Mongolia and northwestern China [5]. An isolated part of the species’ range exists in Transcaucasia [6]. Recently, numerous observations of the species have been reported along the eastern bank of the Ural River [7], which is considered the westernmost limit of its distribution [8].
Data on genetic diversity for this species are scarce and are mostly limited to eastern China [9,10,11]. There are no data available for the northeastern Caspian region, which is the species’ type locality [12].
This study reports the most northerly observations of P. lineolatus west of the Ural River and provides the first data on the species’ mitochondrial DNA diversity in the northeastern Caspian region.

2. Materials and Methods

From 2022 to 2025, we carried out herpetofaunal surveys in the northeastern Caspian area of western Kazakhstan. We searched for snakes through active searches, looking under rocks and flat objects, as well as monitored roads. The exact geographical coordinates of the presence sites were recorded using a GPS tracker (Garmin eTrex H, Garmin Ltd., New Taipei City, Taiwan). We collected tissue samples from encountered snakes, consisting of ethanol-preserved ventral scale clips from live individuals and muscles from road-killed individuals. Alive snakes were released at the place of capture immediately after tissue sampling. Some of the road-killed snakes were collected, and voucher specimens (preserved in 75% alcohol) are deposited in the Herpetological Collection of the M. Utemisov West Kazakhstan University.
The morphological characteristics of a better-preserved road-killed specimen were taken as follows: snout–vent length (SVL) and tail length (both were measured with a tape ruler to the nearest of 1 mm); head length, head width, and head height (taken with a digital caliper to the nearest of 0.1 mm); and number of scale rows at the midbody and number of internasal, prefrontal, frontal, parietal, preocular, postocular, loreal, supralabial, infralabial, temporal, ventral and subcaudal scales. Paired scales on the head were counted on both sides.
Total DNA of P. lineolatus was extracted by a standard procedure including treatment with sodium dodecyl sulfate and proteinase K and subsequent phenol–chloroform extraction [13]. We amplified a standard vertebrate DNA barcode fragment of the mitochondrial gene subunit I of cytochrome c oxidase (COI) using the vertebrate universal primer pair VUTF (5′-TGTAAAACGACGGCCAGTTCTCAACCAAYCAYAARGAYATYGG-3′) and VUTR (5′-CAGGAAACAGCTATGACTARACTTCTGGRTGKCCRAARAAYCA-3′) [14]. The PCR protocol consisted of a 3 min denaturation step at 95 °C, followed by 35 cycles of denaturation for 30 s at 95 °C, primer annealing for 30 s at 54 °C, and extension for 1 min at 72 °C, with a final extension step at 72 °C for 5 min. PCR products were purified by extraction from polyacrylamide gel. Sequencing was performed by the Evrogen company (Moscow, Russia) on an ABI 3500 automated capillary sequencer (Applied Biosystems, Austin, TX, USA) with the BigDye™ Terminator v.3.1 cycle Sequencing Kit (Thermo Fisher Scientific, Vilnius, Lithuania) using the same primers.
The newly obtained sequences were inspected for errors using Chromas 2.6.2 (Technelysium Pty Ltd., Tewantin, Australia) and checked for unexpected stop codons with SeaView 4.4.2 [15]. The sequences were combined with those available in GenBank and manually aligned, resulting in a dataset with a length of 551 bp and 13 sequences. MEGA11 [16] was used to calculate percentage differences (p-distances) between obtained sequences. A haplotype network was generated using the TCS algorithm [17] implemented in Hapsolutely 0.2.3 [18].
We collated the dataset of reliable distribution records (i.e., undoubtful species determination and georeferencing) of the P. lineolatus in regions of western Kazakhstan to comprehensively outline the northwestern distribution limits of the species. Original field survey data, bibliographic records [7,8,19,20,21,22], and research-grade iNaturalist observations (https://www.inaturalist.org, accessed on 22 July 2025) were used to arrange the dataset following the Darwin Core format for biodiversity data. QGIS Desktop 3.30.0 [23] was utilized to create a map and visualize the records.

3. Results

We encountered a total of 30 individuals of P. lineolatus in the West Kazakhstan, Atyrau, and Mangistau regions. The dataset of literature and our new occurrence records for western Kazakhstan totals 104 observations (see Figure 1A and Supplementary Table S1). On 21 April 2025, two road-killed specimens of P. lineolatus were found on the A-28 road between the settlements Tompak and Shabdarjap, west of the Ural River (see Figure 1A,B, localities 1 and 2; Akzhaik District, West Kazakhstan administrative region of Kazakhstan). The specimens were encountered within a 3 km radius of each other (48.8972° N 51.7849° E and 48.8688° N 51.7813° E). The better-preserved adult male specimen from locality 1 was placed into the Herpetological Collection of the M. Utemisov West Kazakhstan University (voucher number ГKЗKУ-44; Figure 2). It has the following morphological characteristics that fit within the known limits for the species: SVL 573 mm, tail length 196 mm, head length 14.9 mm, head width 9.4, head height 4.9; dorsal scales 17, internasals 1/1 (left/right), pre-frontals 1/1, frontal 1, parietal 1/1, preocular, postocular 2/2, loreal 1/1, supralabials 9/9 (4th to 6th supralabials in contact with eyes), infralabials 9/9, temporals 2+3/2+3, ventrals 201, and subcaudal scales were impossible to count due to damages.
On 6 August 2025, another road-killed individual was found in the West Kazakhstan Region (49.5484° N, 53.4218° E), representing the most northern distribution record for the species within its entire geographical range. This specimen was also placed into the collection under voucher number ГKЗKУ-51.
Five distinct COI haplotypes were obtained from ten P. lineolatus individuals from Kazakhstan. New sequences have been deposited in GenBank (NCBI) under accession numbers PX120872-PX120881 (Table 1). The overall average p-distance was 0.81% (maximum 1.63%; Supplementary Table S2). The haplotype network revealed significant structuring (Figure 1C). Individuals from localities 2–4 along the Ural River shared an identical haplotype, while the specimen from locality 1 differed by five substitutions (0.73% p-distance). The same haplotype was also found in the Transcaspian area, on the border between the Atyrau and Mangistau regions of Kazakhstan (locality 5), and in China. Samples from the Ustyurt Plateau (Mangistau Region) formed a separate haplogroup differing by 0.93–1.63%.

4. Discussion

Our findings confirm, for the first time, the distribution of P. lineolatus west of the Ural River, thereby suggesting the species for inclusion in the species list of the European herpetofauna. These localities also represent the northwesternmost records for the species and constitute the first documented observation in the West Kazakhstan administrative region, along with the northernmost observation of the species east of the Ural River. The genus Psammophis has been previously reported from Europe as a fossil, represented by the extinct species P. odysseus Georgalis & Szyndlar, 2022, from the latest Miocene of Salobrena, in Spain [25]. This single occurrence apparently represented a dispersal from northern Africa to the Iberian Peninsula during the Messinian Salinity Crisis. The fact that P. odysseus was found only in that locality implies that Psammophis was short-lived in Europe and that the genus shortly after became extirpated from the continent [25].
The number of records of P. lineolatus from the northwestern distribution limit has increased over the last 25 years. In the review of the distribution and ecology of P. lineolatus in Kazakhstan, Brushko and Kubykin [19] mention that “it barely reaches 48° north latitude,” referring to observations of the species in the Irgiz district of the Aktobe Region by Sushkin at the end of the 19th century and by Kubykin in 1983. However, data obtained in recent years have changed established ideas about the northern limit of the species’ distribution, shifting it west from the Aktobe Region to the Atyrau Region. In 2011, Pestov and co-authors [21] reported the discovery of a steppe ribbon racer in the Inder district of the Atyrau Region (48.1° N). Lately, Saraev (2017) reported numerous observations of the species along the eastern bank of the Ural River, extending as far north as 48.6° N [7]. Further studies in the area revealed the presence of the ribbon racer even farther north (48.8° N) [8]. We observed the species 70 km north of the previous limit (49.5° N). This suggests that species expansion may have occurred during the past few decades rather than being overlooked. Saraev also predicted that P. lineolatus might eventually be observed west of the Ural River, proposing that snakes should use road bridges to cross. The width of the Ural River in the area under discussion (Figure 1B) is relatively small and rarely exceeds 150 m. However, during the flood period, it reaches several kilometers. Furthermore, the occurrence of flooding corresponds to the alleged period of maximal snake activity in April–May. Consequently, the floodplain of the Ural River may act as a substantial, albeit traversable, barrier to P. lineolatus dispersal. Future monitoring studies of this distribution expansion are recommended to better understand the nature of this process, in conjunction with species distribution modeling.
The DNA barcoding survey of the collected P. lineolatus samples yielded pronounced phylogeographical structuring. Notably, two relatively distant haplotypes were found in northwesternmost localities 1 and 2 (Figure 1A,B). One haplotype was also found in an adjacent area along the Ural River; the other seems geographically widespread, having been reported in China, including at the southeasternmost limit of the species’ distribution [9]. This haplotype was also observed in western Kazakhstan, 350 km southeast of the aforementioned location. Although the limited COI data prevents detailed interpretation, these results indicate substantial genetic variation. To date, the most comprehensive study of the steppe ribbon racer intraspecific genetic diversity has focused on China and used the NADH dehydrogenase subunit 4 (ND4) and cytochrome b (CYTB) mtDNA markers [11]. This study resulted in the description of a new species, P. turpanensis Chen, Liu, Cai, Li, Wu, & Guo, 2021. Although the type locality of P. lineolatus was restricted to the Mangystau Region of Kazakhstan [12], no genetic material from this particular area has yet been examined. Our findings suggest that the high degree of genetic variability observed in P. lineolatus in the northwestern Caspian region may lead to intriguing discoveries in future investigations of the species’ phylogeography across its entire range. These studies should incorporate more genetic markers (ND4, CYTB) to integrate along with the previously published data.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/taxonomy5040054/s1, Table S1: Dataset of distribution records of the Psammophis lineolatus in the northeastern Caspian region (western Kazakhstan) (Darwin Core standard). References [26,27,28,29,30,31,32,33] are cited in the Supplementary Materials. Table S2: Uncorrected genetic distances (p-distance; in %) between the examined Psammophis lineolatus specimens, based on COI (551 bp).

Author Contributions

Conceptualization, E.S.; methodology, E.S., A.B. and K.A.; validation, E.S., A.B., A.K. and K.A.; formal analysis, E.S., A.B., A.K., S.L. and K.A.; investigations, E.S., A.B., A.K. and K.A.; resources, E.S., S.L. and K.A.; data curation, E.S., A.B. and A.K.; writing—original draft preparation, K.A. and E.S.; writing—review and editing, E.S., A.B., A.K., S.L. and K.A.; visualization, E.S.; supervision, E.S. and K.A.; project administration, E.S. and K.A.; funding acquisition, K.A. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Science Committee of the Ministry of Science and Higher Education of the Republic of Kazakhstan (grant No. AP 19675960).

Institutional Review Board Statement

The animal study protocol was approved by the Ethics Council of M. Utemisov West Kazakhstan University (No 2480; date of approval 4 September 2023).

Data Availability Statement

All sequences generated in this study were deposited to the GenBank under the accession numbers PX120872-PX120881.

Acknowledgments

We thank all colleagues who provided their occurrence records. The authors also wish to express their gratitude to our field trip driver, Alim Khaidarov, for his instant support during the field work.

Conflicts of Interest

The authors declare no conflicts of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

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Taxonomy 05 00054 g001
Figure 1. Distribution records, studied localities and haplotype network analysis for Psammophis lineolatus in the northeastern Caspian region (western Kazakhstan). (A) The map showing analyzed records of P. lineolatus in Kazakhstan (see legend for details). DNA sampling localities are numbered (refer to Table 1). The administrative boundaries between countries (solid black line) and the boundaries of administrative regions within Kazakhstan (dashed black lines) are shown (AK—Aktobe Region, AT—Atyrau Region; MN—Mangistau Region; WK—West Kazakhstan Region). (B) Detailed map of P. lineolatus observations in the area around the Ural River. Map from Google Maps APIs (2025), NASA, used according to Google Maps APIs Terms of Service. (C) The haplotype network for P. lioneolatus based on COI sequences (551 bp long). The numbers of transverse strokes on the branches correspond to the number of nucleotide substitutions. The numbers on haplotypes correspond to the localities on (A,B). The inset map shows location of analyzed samples within the entire geographical range of P. lineolatus (diagonal fill pattern; according to [24]).
Figure 1. Distribution records, studied localities and haplotype network analysis for Psammophis lineolatus in the northeastern Caspian region (western Kazakhstan). (A) The map showing analyzed records of P. lineolatus in Kazakhstan (see legend for details). DNA sampling localities are numbered (refer to Table 1). The administrative boundaries between countries (solid black line) and the boundaries of administrative regions within Kazakhstan (dashed black lines) are shown (AK—Aktobe Region, AT—Atyrau Region; MN—Mangistau Region; WK—West Kazakhstan Region). (B) Detailed map of P. lineolatus observations in the area around the Ural River. Map from Google Maps APIs (2025), NASA, used according to Google Maps APIs Terms of Service. (C) The haplotype network for P. lioneolatus based on COI sequences (551 bp long). The numbers of transverse strokes on the branches correspond to the number of nucleotide substitutions. The numbers on haplotypes correspond to the localities on (A,B). The inset map shows location of analyzed samples within the entire geographical range of P. lineolatus (diagonal fill pattern; according to [24]).
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Figure 2. Specimen of an adult male Psammophis lineolatus collected west of the Ural River (locality 1, Figure 1A,B) and deposited into the Herpetological Collection of the M. Utemisov West Kazakhstan University (voucher number ГKЗKУ-44, sample ID KZ434, GenBank accession number PX120880).
Figure 2. Specimen of an adult male Psammophis lineolatus collected west of the Ural River (locality 1, Figure 1A,B) and deposited into the Herpetological Collection of the M. Utemisov West Kazakhstan University (voucher number ГKЗKУ-44, sample ID KZ434, GenBank accession number PX120880).
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Table 1. Psammophis lineolatus samples used for sequencing and analysis of COI variability.
Table 1. Psammophis lineolatus samples used for sequencing and analysis of COI variability.
Locality IDLocalityLatitudeLongitudeGenBank Accession Number/Sample IDReference
1WK, Kazakhstan48.897251.7849PX120880/KZ434 *This study
2WK, Kazakhstan48.868851.7813PX120881/KZ436This study
3North shore of lake Inder, AT, Kazakhstan48.512851.9456PX120873/KZ100This study
4Karabatano train station, AT, Kazakhstan47.297252.3098PX120872/KZ32This study
510 km N of Boranqul, AT, Kazakhstan46.310354.3846PX120874/KZ188This study
6Ustyurt National Reserve, MN, Kazakhstan42.563454.2603PX120875/KZ227This study
7Ustyurt National Reserve, MN, Kazakhstan42.570154.2629PX120876/KZ240This study
8Ustyurt National Reserve, MN, Kazakhstan42.595454.2965PX120877/KZ245This study
9Ustyurt National Reserve, MN, Kazakhstan42.595054.2992PX120878/KZ246This study
10Ustyurt National Reserve, MN, Kazakhstan42.597254.2994PX120879/KZ249This study
-Helan Shan, Ningxia, China~38.0~106.0OQ416684/KIZ YPX19277[9]
-Luntai, Xinjiang, China~42.0~84.5OQ416685/KIZ 015468[9]
-Buerjin County, Xinjiang, China47.559387.0565MT991050/GXG311[10]
* Corresponds to voucher number ГKЗKУ-44 (Herpetological Collection of the M. Utemisov West Kazakhstan University). AT—Atyrau Region; MN—Mangistau Region; WK—West Kazakhstan Region.
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MDPI and ACS Style

Akhmedenov, K.; Bakiev, A.; Klenina, A.; Lukonina, S.; Simonov, E. First European Occurrence and Genetic Diversity of the Steppe Ribbon Racer Psammophis lineolatus (Brandt, 1838) (Serpentes: Psammophiidae). Taxonomy 2025, 5, 54. https://doi.org/10.3390/taxonomy5040054

AMA Style

Akhmedenov K, Bakiev A, Klenina A, Lukonina S, Simonov E. First European Occurrence and Genetic Diversity of the Steppe Ribbon Racer Psammophis lineolatus (Brandt, 1838) (Serpentes: Psammophiidae). Taxonomy. 2025; 5(4):54. https://doi.org/10.3390/taxonomy5040054

Chicago/Turabian Style

Akhmedenov, Kazhmurat, Andrey Bakiev, Anastasia Klenina, Svetlana Lukonina, and Evgeniy Simonov. 2025. "First European Occurrence and Genetic Diversity of the Steppe Ribbon Racer Psammophis lineolatus (Brandt, 1838) (Serpentes: Psammophiidae)" Taxonomy 5, no. 4: 54. https://doi.org/10.3390/taxonomy5040054

APA Style

Akhmedenov, K., Bakiev, A., Klenina, A., Lukonina, S., & Simonov, E. (2025). First European Occurrence and Genetic Diversity of the Steppe Ribbon Racer Psammophis lineolatus (Brandt, 1838) (Serpentes: Psammophiidae). Taxonomy, 5(4), 54. https://doi.org/10.3390/taxonomy5040054

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