Phylogeny and geographic distribution of rock lizards (Lacertidae, Reptilia) in Alborz mountain range

Cover Page

Cite item

Full Text

Abstract

Rock lizards of the genus Darevskia have long been an important model object for study of reptile evolution. To understand the overall picture, it is important to know how bisexual and parthenogenetic species are distributed within this genus. The ranges of Caucasian species of rock lizards have been studied for a long time and in detail. However, recent attention to the species inhabiting the territory of Iran has been attracted after the description of several new species in 2013. As part of the continuation of these studies, we studied the distribution and genetic diversity of five species of lizards of the Darevskia genus along the Alborz mountain range in Iran: D. chlorogaster, D. caspica, D. defilippii, D. schaekeli and D. raddei. In the course of this, we discovered new localities of D. defilippii that are outside their known range. We assessed the phylogenetic relationships between the studied species according to the ND4 – Leu tRNA region of mitochondrial DNA and found a relatively high level of genetic variability in D. defilippii and D. chlorogaster, while D. raddei, despite its wide distribution, has low variability. In general, the phylogenetic position of the studied species is somewhat different from that described by Ahmadzadeh et al. (2013).

Full Text

Introduction

Rock lizards of Darevskia genus comprise about 40 species including seven parthenogenetic species. Most of them are distributed on the Armenian Highland (Darevky, 1967), however, some species occupy rocky areas spreading to Turkmenistan in the East, and Eastern Europe, Balkans and Turkey in the West. Only a few works on rock lizards were devoted to distribution and phylogeny of rock lizards in Iran that occupied Alborz and Kopet Dagh mountain ranges (Ahmadzadeh et al., 2013; Rastegar- Pouyani et al., 2013; Yousefi et al., 2013). Separation of two new species from D. chlorogaster complex (D. caspica, D. kamii) and two species from D. defilippii complex (D. kopetdaghica, D. schaekeli) in Alborz mountains by Ahmadzadeh et al. (2013) demonstrates that taxonomy and distribution of rock lizards from this rich region were not studied well enough. Thus, we set the aim to check the relevance of distribution areas and estimate phylogenetic position of rock lizard species inhabiting western and central Alborz mountains after nine years have passed since discovery of the new species.

Species ranges can change quite strongly and quickly within a short time for many reasons. Ranges of reptiles and amphibians may shrink or expand due to the direct anthropogenic influence, such as habitat modification (Doherty et al., 2020), noise (Simmons, Narins, 2018) and probably light pollution (Perry et al., 2008), and indirect influence, for example climate change (Erasmus et al., 2002; Peck et al., 2009). In this case, there are difficulties associated with preventing the extinction of species and climate change observation.

Alborz mountain range is one of the main centres of origin of rock lizards (Ahmadzadeh et al., 2013; Freitas et al., 2016). Due to the diversity of the landscape and, consequently, the variety of biotopes, geographic and ecological isolation is formed between species and populations within them. In this regard, the study of the genetic diversity and distribution of rock lizards on Alborz will expand the knowledge of the evolution of the Darevskia genus.

Material and methods

Sampling and Mapping. We collected the material along the western and central parts of Alborz mountain range along the southern coast of the Caspian Sea in Iran from the spring-summer period of 2022. During the expedition we carefully searched for lizards along the roads and four times crossed the mountain ridge from North to South and vice versa in different parts; we were also guided by the distribution map in the work of Ahmadzadeh et al. (2013). Visually detected lizards were captured by noose or by hand; dorsal and ventral pictures of each lizard were taken by telephone photo camera (IPhone 12 mini). Iryshkov, Arakelyan, Moaddab, Milto and Galoyan participated in the capture of lizards.

Tail tips of the lizards were fixed in 96% ethanol. Following species were included: 3 sp. belonging to D. caspica, 13 sp. to D. chlorogaster, 10 sp. to D. defilippii, 41 sp. to D. raddei and 4 sp. to D. schaekeli. Samples are stored in two instances: at the Hakim Sabzevari University and at the Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences.

We took GPS coordinates of each finding using GPS tracker (Garmin 64S, WGS 84) and put the point on the map. Also we added terrestrial ecoregions according to bioregions word map from oneearth.org (2001). The basis of species distribution map was taken from Wikimedia Commons author NordNordWest, licensed under CC BY- SA 3.0; colours were changed, labels, points, ecoregions, scale bar and north arrow were added for the article.

When determining the distance between the capture points and the range boundary of D. defilippii, we calculated the distance between our point and the northwesternmost point presented in Ahmadzadeh et al. (2013) using Google maps. Then we rounded the distance up to integer values of kilometres. Using Google maps, we also obtained the approximate height of each GPS point presented in the supplementary material from Ahmadzadeh et al., 2013 and added altitudes measured by us (presented in Table 1). We rounded the resulting height values up to tens.

DNA Extraction, Amplification and Sequencing. We isolated genomic DNA from the 96% ethanol-fixed tissue samples using the SDS (Sodium Dodecyl Sulfate) DNAextraction method (Kabir et al., 2006), and the quality of extracted DNAwas measured using NanoDrop 1000. We chose the mitochondrial DNA fragment ND4 – Leu tRNA (907 bp) which includes fragment of the NADH dehydrogenase 4 and His, Ser and Leu tRNAs (Arevalo et al., 1994) due to its sufficient variability and the presence of a large number of reptile sequences in the GenBank.

For polymerase chain reaction (PCR) we used ND4 (forward) and Leu (reverse) primers and stuck to the amplification protocol described by Dutton et al. (1996). PCR products were sequenced by the Microsynth DNA company using the forward primer and the manufacture protocols.

Phylogenetic Analysis. The sequences were aligned in the program MEGA11 and adjusted manually. We used the sequences of Podarcis muralis [Laurenti, 1768] and P. siculus [Rafinesque-Schmaltz, 1810] (NC_011607.1; NC_011609.1) from the GenBank as an outgroup in alignment.

We used two types of datasets for phylogenetic analysis. First was a reduced dataset, including only our sequences and outgroups from GenBank, and second was a full dataset, including our sequences and sequences from the work of Ahmadzadeh et al. (2013) with outgroups from GenBank.

Phylogenetic trees were reconstructed under Bayesian criteria (BI) and using the maximum likelihood (ML) method. The optimal partitioning schemes and models for Bayesian Inference analysis were identified with PartitionFinder software (Lanfear et al., 2012) using greedy search algorithm under AIC criterion. Following schemes were identified: F81 for 1st position of ND4, GTR+G for 2d and 3d positions separately, and HKY+G for tRNAs (for reduced dataset); for the full dataset HKY+G for 1st position of ND4, GTR+G for 2d, HKY+I+G for 3d position and for tRNAs partition separately (for full dataset). BI was performed using MrBayes v3.2.6 (Ronquist, Huelsenbeck, 2003) with two simultaneous runs, each with four chains, for 5 million generations. We checked the convergence of the runs and that the effective sample sizes (ESS) were all above 200 by exploring the likelihood plots using TRACER v1.7.1 (Rambaut et al., 2018). The initial 10% of trees were discarded as burnin. Confidence in tree topology was assessed by posterior probability (PP) (Huelsenbeck, Ronquist, 2001). The ML trees were generated using IQtree software (Nguyen et al., 2015) with ultrafast bootstrap = 10,000 (UFBoot) (Minh et al., 2013), partitioning schemes and models were selected using ModelFinder software (Kalyaanamoorthy et al., 2017). Following schemes were selected: F81+F for 1st position of ND4, TN+F for 2d, TN+F+G4 for 3d position and separately for tRNAs partition (for full dataset); F81+F for 1st position of ND4, TN+F+G4 for 2d, HKY+F+G4 for 3d position and separately for tRNAs partition (for full dataset).

For visualisation we used dendrograms according to the Bayesian method (BI) and also added the values obtained by maximum likelihood (ML) analysis. The dendrograms were processed using FigTree v1.4.4 (tree.bio.ed.ac.uk/ software/figtree/). We calculated the uncorrected interand intragroup pairwise distances (p-distances) in the MEGA11 program using 1000 bootstrap replicas.

 

Table 1. Specimens used for phylogenetic analysis with localities and GenBank numbers of ND4 – Leu sequences obtained by us

 

Results and discussion

The topologies of the phylogenetic tree of our sequences, obtained by the BI and ML methods, are consistent in all nodes, but somewhat different in the levels of support. Phylogenetic relationships between Chlorogaster (includes D. chlorogaster and D. caspica) and Defilippii (includes D. defilippii and D. schaekeli) clades has temperate support – 0.94/79. D. chlorogaster and D. caspica are separated with high support, while the node between D. defilippii and D. schaekeli has a high support only by BI  (Fig. 1).

According to Figure 1 and Table 2 D. chlorogaster and D. caspica are genetically close, as they belong to the same clade and p-distance between them is 6.22%. D. schaekeli and D. defilippii are also closely related, with p-distance of 9.02% since they belong to the same clade. Distances between species from different clades are much greater: 10.84% between D. chlorogaster and D. defilippii, 11.59% between D. chlorogaster and D. schaekeli, 12.74% between D. caspica and D. defilippii, and 12.69% between D. caspica and D. schaekeli. D. raddei belongs to the other clade – Raddei, it has a large distances to other species and occupies basal position on trees topology (Figs. 1, 2). Generally, D. raddei is a monophyletic group (1/98 support in Fig. 1) with a low genetic variability (Table 2). It coincides with the results published in Freitas et al. (2016).

During visual sequence check we noticed a possible nuclear mitochondrial pseudogene (Bensasson et al., 2001) of ND4 – Leu tRNAfragment – the external parts of sequences were not completely read and a large number of double peaks on the electropherogram were observed (Chow et al., 2021). This was not noticed or reported in the closely related species D. caspica and other distant species.

Unresolved nodes (Fig. 1) in D. caspica and D. schaekeli together with their low within-group p-distances indicate low genetic variability within these species  (Table 2).

As shown in the dendrogram reconstructed from our sequences (Fig. 1), D. defilippii indeed has a fairly high genetic variability. This might be explained by the diverse habitats inhabited by D. defilippii (Ahmadzadeh et al., 2013). Phylogenetic relationships within D. defilippii reflect its geographical structure. Lineage consisting of specimens from Halu Dasht and Nilu villages from Gilan province diverged with the high level of support by BI, but with low support by ML – 1/88. Specimen from Vadarbon village, Mazandaran province (OR340751) separates furthest with high node support – 1/99. There is no connection between phylogenetic structure of D. chlorogaster and its geographic dispersal as in D. defilippii, although D. chlorogaster is also represented in several localities in Gilan province. However, genetic variability (based on uncorrected p-distances) within D. chlorogaster is two times stronger than in D. defilippii (2 and 1% respectively) (Table 2).

 

Fig. 1. Dendrogram showing phylogenetic relationships between rock lizards in different regions of Iran, based on the sequences of the ND4 – Leu tRNA fragment. Values over and under the nodes represent posterior probabilities by BI/bootstrap values by ML. The localities for each specimen are shown to the right of the vertical thin black line

 

Table 2. Uncorrected p-distances (%) of reduced and full datasets of ND4 – Leu mDNA fragment: Intergroup distances are shown under the diagonal, intragroup distances – on the diagonal, standard errors – above the diagonal

 

Species

D. chlorogaster

D. caspica

D. schaekeli

D. defilippii

D. raddei

D. chlorogaster

2

0.88

1.16

1.06

0.91

D. caspica

6.22

0

1.23

1.1

1.1

D. schaekeli

11.59

12.69

0

0.98

1.16

D. defilippii

10.84

12.74

9.02

1

0.97

D. raddei

11.59

12.69

13.13

10.66

0

 

It contradicts the results obtained in Ahmadzadeh et al. (2013), where analysis of cytochrome b and ND4 genes concluded that the variability of this species is low. In this regard, it can be assumed that the genetic variability of D. defilippii is primarily exists due to geographic isolation in populations, while the causes of genetic diversity in D. chlorogaster are obscure.

With the addition of sequences from hmadzadeh et al. (2013) the topology of the tree changed (Fig. 2). Relationships between main clades and between D. defilippii and D. schaekeli are resolved with a high support unlike the first tree, that contains only our sequences (Fig. 1). D. caspica is not monophyletic, it splits into three separate clusters. One cluster has high support, while two others form a trichotomy with D. chlorogaster within a lowsupported group (0.63/70). D. schaekeli specimens with high levels of support are divided according to localities from different provinces. Localities from Mazandaran province are divided separately – with individuals caught by us in Darreh-ye Malek Darreh and individuals from GenBank from Savasraeh (KF717317) and Kiasar (KF717316) which are the most distant from the others. D. defilippii specimens form a clade with trichotomy divided by localities Eshkevarat, Halu Dasht, and Nilu villages from Gilan province. Phylogenetic distribution of D. defilippii by localities has not been confirmed: nodes do not form distinct clusters and nodes supports are low. Phylogenetic relationships within D. chlorogaster still remain unclear: individual clusters by locality have not been formed, almost all nodes have low support, and many of them are unresolved. However, the most distant specimen within D. chlorogaster clade is from Dohezar, Mazandaran province (KF717320), and it is significantly divided from others.

 

Fig. 2. Dendrogram showing phylogenetic relationships between rock lizards in different regions of Iran and some localities from Azerbaijan based on sequences of the ND4 – Leu tRNAfragment obtained by us and sequences of ND4 gene by Ahmadzadeh et al. (2013) from GenBank. Values over and under the nodes represent posterior probabilities by BI/bootstrap values by ML. Branches coloured by species according to Figure 1. Blue coloured sequence numbers – received by us, black – by Ahmadzadeh at al. (2013). The localities for each specimen are shown to the right of the vertical thin black line

 

We have also discovered two new locality records of D. defilippii beyond the distribution range (Ahmadzadeh et al., 2013). Records were taken 23 (OR340778 and OR340780 sequences) and 30 km (OR340724 and OR340725 sequences) northwest of previously documented locations. On the first point we caught individuals on a dry sandstone slope along an unpaved road surrounded by steppe and on the second point on a rocky slope with cracks along an asphalt road surrounded by mixed forest. In this regard, it can be concluded that either the range is initially wider, or it could expand towards the northwest during the last nine years.

The presented species also have different altitude distributions. D. caspica occupies Hyrcanian mixed forest (Fig. 3) at an altitude mainly between 130 and 580 metres a.s.l., but also there was a population from the altitude of about 1590 m in Savasraeh (KF717318 sequence). Although it separated from the specimens (KF717316, KF717317 sequences) from Joybar (altitude is about 400 m a.s.l.), it forms with these specimens one group with 0.99/97 support (Fig. 2) which indicates the lack of strong isolation.

D. chlorogaster has a range with a large elevation difference: from 10 to 1250 m a.s.l., and confined to Hyrcanian forest and forest-steppe ecozones. D. raddei also occupies a wide elevation range of 840–2040 m a.s.l. within forest and forest-steppe zones. D. defilippii lives in the high altitudes of 1290–2500 m a.s.l. in the forest-steppe. Moreover, an altitudinal distribution of this species is wider – some populations are known from the forested areas at 440 and 640 m a.s.l. D. schaekeli is also a high altitudinal species and, according to our and already published data, distributed only at 1560–2300 m a.s.l. in forest-steppe ecozones.

The Hyrcanian forest consists of a large variety of woody plant species with high densities. The humidity level is high due to the large number of mountain streams flowing down the mountain slopes (Naqinezhad et al., 2008; Jafari et al., 2013), high precipitation (Heshmati, 2007) and mild climate due to low altitude (Beck et al., 2018). The foreststeppes are characterised by their mosaic vegetation: in some areas there are quite dense wet forest fragments and arid steppe spaces between them. Precipitation is generally low (Heshmati, 2007). Due to the high altitude, there is a high annual temperature range (Beck et al., 2018). Desert basins are characterised by sparse vegetation and low precipitation. Daily and annual temperature differences there are quite high (Breckle, 2002). In this regard, it can be concluded that the existing distribution of species can be explained not only by altitude, but also by preferred humidity level and vegetation. Available data of distribution suggest that D. chlorogaster and D. caspica prefer wet habitats with dense vegetation, while D. defilippii and D. schaekeli prefer dry, sparsely vegetated but more rocky biotopes. D. raddei, apparently, can occupy an intermediate position between the presented species, but at the moment there is not enough data for accurate conclusions.

 

Fig. 3. Distribution of studied species of rock lizards within Alborz mountain ridge in Iran. Small circles indicate localities of the species according to Ahmadzadeh et al. (2013). Big circles indicate the localities we discovered during the expedition. Each colour corresponds to the species of lizards and coincides with the colour from Figs. 1, 2. Coloured areas – terrestrial ecoregions according to Safaei-Mahroo et al. (2015) that showed only for Iran

 

It is important to note that the map presented in Safaei-Mahroo et al. (2015) was built using data of Olson et al. obtained in 2001. Therefore, the distribution of ecoregions at the moment of our data collection (2022) can be significantly different due to the desertification (Cherlet et al., 2018). In particular, the localities we found (Fig. 3) are located in the forest zone, but this zone, according to our observations, is a forest-steppe. For this reason, the range of D. defilippii could be shifted to the northwest.

Although studied species have some differences in their preferred biotopes, the borders between them are very fuzzy. Thus, further research should be aimed at clarifying the boundaries of the ranges of species and studying the reasons for such distribution.

Conclusions

  1. We confirmed that the species defilippii has a high genetic variability mediated by geographic isolation.
  2. chlorogaster is one of the most genetically variable among studied species. This may be explained by the possible presence of a nuclear mitochondrial ND4 – Leu tRNA fragment pseudogene.
  3. Despite the wide distribution range and diverse habitat preferences raddei has a low variability of the ND4 – Leu tRNA fragment of mDNA.
  4. The range of the species defilippii is broader than previously suggested.
  5. The distribution limits for considered species require clarification and presence of the secondary contact zones is very

Acknowledgments

We thank Andrey Stukalov and Irena Martirosyan for help in interpreting results of this work, Ali Moaddab for assistance in organising transportation to and within the study areas and Victor Spangenberg for participation in the collection and transportation of the material.

×

About the authors

Eugene S. Iryshkov

Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences; Lomonosov Moscow State University

Author for correspondence.
Email: eiryshkov@gmail.com
ORCID iD: 0000-0003-4171-8396
Russian Federation, 33 Leninsky Prosp., Moscow 119071; 1 Vorobyovi Gory, Moscow 119992

Eugenia N. Solovyeva

Lomonosov Moscow State University

Email: anolis@yandex.ru
ORCID iD: 0000-0001-7564-9187

Zoological Museum

Russian Federation, 2 Bolshaya Nikitskaya St., Moscow 125009

Marine S. Arakelyan

Yerevan State University

Email: arakelyanmarine@gmail.com
ORCID iD: 0000-0002-6334-5714
Armenia, 1 Alex Manoogian St., Yerevan 0025

Eskandar Rastegar-Poyani

Hakim Sabzevari University

Email: rastegarpouyani45@gmail.com
ORCID iD: 0000-0002-9639-2058
Iran, Islamic Republic of, 8 Tovhid, Sabzevar 9617976487

Marteza Moaddab

Hakim Sabzevari University

Email: rastegarpouyani45@gmail.com
Iran, Islamic Republic of, 8 Tovhid, Sabzevar 9617976487

Konstantin D. Milto

Zoological Institute of the Russian Academy of Sciences

Email: coluber@zin.ru
ORCID iD: 0000-0003-4061-0164
Russian Federation, 1 Universitetskaya embankment, St. Petersburg 199034

Eduard A. Galoyan

Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences; Lomonosov Moscow State University

Email: saxicola@mail.ru
ORCID iD: 0000-0001-7633-7759

Zoological Museum

Russian Federation, 33 Leninsky Prosp., Moscow 119071; 2 Bolshaya Nikitskaya St., Moscow 125009

References

  1. Ahmadzadeh F., Flecks M., Carretero M. A., Mozaffari O., Böhme W., Harris D. J., Freitas S., Rödder D. Cryptic speciation patterns in Iranian rock lizards uncovered by integrative taxonomy. PLoS ONE, 2013, vol. 8, no. 12, article no. e80563. https://doi.org/10.1371/journal.pone.0080563
  2. Arevalo E., Davis S. K., Sites Jr. J. W. Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology, 1994, vol. 43, no. 3, pp. 387–418. https://doi.org/10.1093/sysbio/43.3.387
  3. Beck H. E., Zimmermann N. E., McVicar T. R., Vergopolan N., Berg A., Wood E. F. Present and future köppengeiger climate classification maps at 1-km resolution. Scientific Data, 2018, vol. 5, iss. 1, pp. 1–12. https://doi.org/10.1038/sdata.2018.214
  4. Bensasson D., Zhang D. X., Hartl D. L., Hewitt G. M. Mitochondrial pseudogenes: Evolution’s misplaced witnesses. Trends in Ecology and Evolution, 2001, vol. 16, iss. 6, pp. 314–321. https://doi.org/10.1016/S0169-5347(01)02151-6
  5. Breckle S. W. Salt deserts in Iran and Afghanistan. Sabkha Ecosystems, 2002, vol. 1, pp. 109–122.
  6. Cherlet M., Hutchinson C., Reynolds J., Hill J., Sommer S., von Maltitz G. World Atlas of Desertification. Luxembourg, Publication Office of the European Union, 2018. 252 p. https://doi.org/10.2760/06292
  7. Chow S., Yanagimoto T., Takeyama H. Detection of heteroplasmy and nuclear mitochondrial pseudogenes in the Japanese spiny lobster Panulirus japonicus. Scientific Reports, 2021, vol. 11, iss. 1, article no. 21780. https://doi.org/10.1038/s41598-021-01346-8
  8. Darevsky I. S. Rock Lizards of the Caucasus: Systematics, Ecology, and Phylogenesis of the Polymorphic Groups of Caucasian Rock Lizards of the Subgenus Archaeolacerta. Leningrad, Nauka, 1967. 214 p. (in Russian).
  9. Doherty T. S., Balouch S., Bell K., Burns T. J., Feldman A., Fist C., Garvey T. F., Jessop T. S., Meiri S., Driscoll D. A. Reptile responses to anthropogenic habitat modification: A global meta-analysis. Global Ecology and Biogeography, 2020, vol. 29, iss. 7, pp. 1265–1279. https://doi.org/10.1111/geb.13091
  10. Dutton P. H., Davis S. K., Guerra T., Owens D. Molecular phylogeny for marine turtles based on sequences of the ND4-leucine tRNA and control regions of mitochondrial DNA. Molecular Phylogenetics and Evolution, 1996, vol. 5, iss. 3, pp. 511–521. https://doi.org/10.1006/mpev.1996.0046
  11. Erasmus B. F. N., Van Jaarsveld A. S., Chown S. L., Kshatriya M., Wessels K. J. Vulnerability of South African animal taxa to climate change. Global Change Biology, 2002, vol. 8, iss. 7, pp. 679–693. https://doi.org/10.1046/j.1365-2486.2002.00502.x
  12. Freitas S., Rocha S., Campos J., Ahmadzadeh F., Corti C., Sillero N., Ilgaz C., Kumlutaş Y., Arakelyan M., Harris D. J., Carretero M. A. Parthenogenesis through the ice ages: A biogeographic analysis of Caucasian rock lizards (genus Darevskia). Molecular Phylogenetics and Evolution, 2016, vol. 102, pp. 117–127. https://doi.org/10.1016/j.ympev.2016.05.035
  13. Heshmati G. A. Vegetation characteristics of four ecological zones of Iran. International Journal of Plant Production, 2007, vol. 1, no. 2, pp. 215–224.
  14. Huelsenbeck J. P., Ronquist F. MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 2001, vol. 17, no. 8, pp. 754–755.
  15. Jafari S. M., Zarre S., Alavipanah S. K. Woody species diversity and forest structure from lowland to montane forest in Hyrcanian forest ecoregion. Journal of Mountain Science, 2013, vol. 10, iss. 4, pp. 609–620. https://doi.org/10.1007/s11629-013-2652-2
  16. Kabir S., Shahriar M., Kabir A. H., Uddin M. G. High salt SDS-based method for the direct extraction of genomic DNA from three different gram-negative organisms. CDR, 2006, vol. 1, no. 2, pp. 57–64.
  17. Kalyaanamoorthy S., Minh B. Q., Wong T. K., Von Haeseler A., Jermiin L. S. ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods, 2017, vol. 14, no. 6, pp. 587–589. https://doi.org/10.1038/nmeth.4285
  18. Lanfear R., Calcott B., Ho S. Y., Guindon S. Partitionfinder: Combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 2012, vol. 29, iss. 6, pp. 1695–701. https://doi.org/10.1093/molbev/mss020
  19. Minh B. Q., Nguyen M. A. T., Von Haeseler A. Ultrafast Approximation for phylogenetic bootstrap. Molecular Biology and Evolution, 2013, vol. 30, iss. 5, pp. 1188–1195. https://doi.org/10.1093/molbev/mst024
  20. Naqinezhad A., Hamzeh’ee B., Attar F. Vegetation–environment relationships in the alderwood communities of Caspian lowlands, N. Iran (toward an ecological classification). Flora: Morphology, Distribution, Functional Ecology of Plants, 2008, vol. 203, iss. 7, pp. 567–577. https://doi.org/10.1016/j.flora.2007.09.007
  21. Nguyen L. T., Schmidt H. A., von Haeseler A., Minh B. Q. IQ-TREE: A Fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution, 2015, vol. 32, iss. 1, pp. 268–274. https://doi.org/10.1093/molbev/msu300
  22. Olson D. M., Dinerstein E., Wikramanayake E. D., Burgess N. D., Powell G. V. N, Underwood E. C., D’Amico J. A., Itoua I., Strand H. E., Morrison J. C., Loucks C. J., Allnutt T. F., Ricketts T. H., Kura Y., Lamoreux J. F., Wettengel W. W., Hedao P., Kassem K. R. Terrestrial ecoregions of the world: A new map of life on Earth. BioScience, 2001, vol. 51, pp. 933–938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
  23. Peck L. S., Clark M. S., Morley S. A., Massey A., Rossetti H. Animal temperature limits and ecological relevance: Effects of size, activity and rates of change. Functional Ecology, 2009, vol. 23, iss. 2, pp. 248–256. https://doi.org/10.1111/j.1365-2435.2008.01537.x
  24. Perry G., Buchanan B. W., Fisher R. N., Salmon M., Wise S. E. Effects of artificial night lighting on amphibians and reptiles in urban environments. Urban Herpetology, 2008, vol. 3, pp. 239–256.
  25. Peterson D. L., Gleisner J. M., Blakley R. L. Bovine liver dihydrofolate reductase. Purification and properties of the enzyme. Biochemistry, 1975, vol. 14, no. 24, pp. 5261–5267. https://doi.org/10.1021/bi00695a005
  26. Rambaut A., Drummond A. J., Xie D., Baele G., Suchard M. A. Posterior Summarization in Bayesian Phylogenetics Using Tracer 1.7. Systematic Biology, 2018, vol. 67, no. 5, pp. 901–904. https://doi.org/10.1093/sysbio/syy032
  27. Rastegar-Pouyani E., Rastegar-Pouyani N., Hosseinian Yousefkhani S. S., Arab M. Rediscovery of Darevskia steineri (Eiselt, 1995) (Sauria: Lacertidae) from Iran. Russian Journal of Herpetology, 2013, vol. 20, no. 1, pp. 36–38. https://doi.org/10.30906/1026-2296-2013-20-1-36-38
  28. Ronquist F., Huelsenbeck J. P. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 2003, vol. 19, no. 12, pp. 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
  29. Safaei-Mahroo B., Ghaffari H., Fahimi H., Yazdanian S., Majd E., Rezazadeh S., Hosseinzadeh M., Nasrabadi R., Rajabizadeh M., Mashayekhi M., Motesharei A., Nader A., Kazemi S. The herpetofauna of Iran: Checklist of taxonomy, distribution and conservation status. Asian Herpetological Research, 2015, vol. 6, no. 4, pp. 257–290. https://doi.org/10.16373/j.cnki.ahr.140062
  30. Simmons A. M., Narins P. M. Effects of anthropogenic noise on amphibians and reptiles. In: Effects of Anthropogenic Noise on Animals. New York, Springer, 2018, vol. 66, pp. 179– 208. https://doi.org/10.1007/978-1-4939-8574-6_7
  31. Yousefi M., Khani A., Eslahi H., Hosseinian Yousefkhani S. S. Easternmost record of Darevskia defilippii (Camerano, 1877) from Qarchaqe Protected Area, Kopet Dagh Mountains, northeastern Iran. L@certidae, 2013, vol. 9, pp. 160–163.

Supplementary files

Supplementary Files
Action
1. JATS XML
Download
2. Table 1. Specimens used for phylogenetic analysis with localities and GenBank numbers of ND4 – Leu sequences obtained by us

Download (770KB)
Indexing metadata
3. Fig. 1. Dendrogram showing phylogenetic relationships between rock lizards in different regions of Iran, based on the sequences of the ND4 – Leu tRNA fragment. Values over and under the nodes represent posterior probabilities by BI/bootstrap values by ML. The localities for each specimen are shown to the right of the vertical thin black line

Download (710KB)
Indexing metadata
4. Fig. 2. Dendrogram showing phylogenetic relationships between rock lizards in different regions of Iran and some localities from Azerbaijan based on sequences of the ND4 – Leu tRNAfragment obtained by us and sequences of ND4 gene by Ahmadzadeh et al. (2013) from GenBank. Values over and under the nodes represent posterior probabilities by BI/bootstrap values by ML. Branches coloured by species according to Figure 1. Blue coloured sequence numbers – received by us, black – by Ahmadzadeh at al. (2013). The localities for each specimen are shown to the right of the vertical thin black line

Download (816KB)
Indexing metadata
5. Fig. 3. Distribution of studied species of rock lizards within Alborz mountain ridge in Iran. Small circles indicate localities of the species according to Ahmadzadeh et al. (2013). Big circles indicate the localities we discovered during the expedition. Each colour corresponds to the species of lizards and coincides with the colour from Figs. 1, 2. Coloured areas – terrestrial ecoregions according to Safaei-Mahroo et al. (2015) that showed only for Iran

Download (810KB)
Indexing metadata

Copyright (c) 2023 Russian Academy of Sciences

Согласие на обработку персональных данных с помощью сервиса «Яндекс.Метрика»

1. Я (далее – «Пользователь» или «Субъект персональных данных»), осуществляя использование сайта https://journals.rcsi.science/ (далее – «Сайт»), подтверждая свою полную дееспособность даю согласие на обработку персональных данных с использованием средств автоматизации Оператору - федеральному государственному бюджетному учреждению «Российский центр научной информации» (РЦНИ), далее – «Оператор», расположенному по адресу: 119991, г. Москва, Ленинский просп., д.32А, со следующими условиями.

2. Категории обрабатываемых данных: файлы «cookies» (куки-файлы). Файлы «cookie» – это небольшой текстовый файл, который веб-сервер может хранить в браузере Пользователя. Данные файлы веб-сервер загружает на устройство Пользователя при посещении им Сайта. При каждом следующем посещении Пользователем Сайта «cookie» файлы отправляются на Сайт Оператора. Данные файлы позволяют Сайту распознавать устройство Пользователя. Содержимое такого файла может как относиться, так и не относиться к персональным данным, в зависимости от того, содержит ли такой файл персональные данные или содержит обезличенные технические данные.

3. Цель обработки персональных данных: анализ пользовательской активности с помощью сервиса «Яндекс.Метрика».

4. Категории субъектов персональных данных: все Пользователи Сайта, которые дали согласие на обработку файлов «cookie».

5. Способы обработки: сбор, запись, систематизация, накопление, хранение, уточнение (обновление, изменение), извлечение, использование, передача (доступ, предоставление), блокирование, удаление, уничтожение персональных данных.

6. Срок обработки и хранения: до получения от Субъекта персональных данных требования о прекращении обработки/отзыва согласия.

7. Способ отзыва: заявление об отзыве в письменном виде путём его направления на адрес электронной почты Оператора: info@rcsi.science или путем письменного обращения по юридическому адресу: 119991, г. Москва, Ленинский просп., д.32А

8. Субъект персональных данных вправе запретить своему оборудованию прием этих данных или ограничить прием этих данных. При отказе от получения таких данных или при ограничении приема данных некоторые функции Сайта могут работать некорректно. Субъект персональных данных обязуется сам настроить свое оборудование таким способом, чтобы оно обеспечивало адекватный его желаниям режим работы и уровень защиты данных файлов «cookie», Оператор не предоставляет технологических и правовых консультаций на темы подобного характера.

9. Порядок уничтожения персональных данных при достижении цели их обработки или при наступлении иных законных оснований определяется Оператором в соответствии с законодательством Российской Федерации.

10. Я согласен/согласна квалифицировать в качестве своей простой электронной подписи под настоящим Согласием и под Политикой обработки персональных данных выполнение мною следующего действия на сайте: https://journals.rcsi.science/ нажатие мною на интерфейсе с текстом: «Сайт использует сервис «Яндекс.Метрика» (который использует файлы «cookie») на элемент с текстом «Принять и продолжить».