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7. Arakelyan M. The study of age, growth, and longevity in tryploid hybrids of rock lizards of genus Darevskia in Armenia // Russian Journal of Herpetology, 2002, 9 1: 63-68.

8. Arakelyan M., Jenderedjan A. Age structure of population of the sand lizards Lacerta agilis in Armenia // 12th Ordinary General Meeting Societias Europaea Herpetologica, Saint-Petersburg, Russia, 12-16 August, 2003: 31-32.

9. Chazaryan L, Vartanyan L., Arakelyan M. New data about helminthes of Colubrid snakes // 12th Ordinary General Meeting Societias Europaea Herpetologica, Saint Petersburg, Russia, 12-16 August, 2003: 49.

10. Krasnov B., Sarfati M., Arakelyan M., Khokhlova I., Burdelova N., Degen A. Host specificity and foraging efficiency in blood-sucking parasite: feeding patterns of the flea Parapulex chephrenison two species of desert rodents // Parasitological Research, 2003, 90: 393399.

11. Arakelyan M. Comparison of parasitic infestation in parthenogenetic and bisexual species of rock lizards // 5th International Symposium on the Lacertids, Lipari, Aeolian Islands, Sicily, Italy, 7-11 May, 2004: 3.

12. Fielden L.J., Krasnov B.R., Khokhlova I.S., Arakelyan M.S. Respiratory gas exchange in the desert flea Xenopsylla ramesis (Siphonaptera: Pulicidae): response to temperature and blood-feeding // Comparative Biochemistry and Physiology, 2004, 137: 557565.

13. Taskavak E., Arakelyan M., Parham J. Testudo graeca armeniaca // Manduria, 2004, 7 (22): 7-8.

14. ., ., . , , Eremias strauchi (Reptilia, Lacertidae) // 1- , , , 10 12 , 2005: 9-12.

15. Krasnov B.R., Khokhlova I.S., Arakelyan M.S., Degen A.A. Is a starving host tastier? Reproduction in fleas parasitizing food-limited rodents // Functional Ecology, 2005, 19: 625631.

16. Parham J., Trkozan O., Stuart B., Arakelyan M., Shafei S., Macey J., Papenfuss T. Genetic evidence for premature taxonomic inflation in Middle Eastern tortoises // Proceedings of the California Academy of Sciences, 2006, 57(3): 955-964.

17. Arakelyan M. The ratio of blood parasites between hybrids and their parental species of rock lizards of genus Darevskia // 14th European Congress of Herpetology, Porto, Portugal, 19-23 September, 2007: 170.

18. Fritz U., Guicking D., Kami H., Arakelyan M., Auer M., Ayaz D., Fernandez C., Bakiev A., Celani A., Duki G., Fahd S., Hava P., Joger U., Khabibullin V., Mazanaeva L., iroky P., Tripep S., Velez A., Anton G., Wink M. Mitochondrial phylogeography of European pond turtles (Emys orbicularis, Emys trinacris) an update // Amphibia-Reptilia 28, 2007: 418-426.

19. Stepanyan I., Arakelyan M., Danielyan F. Tetraploid hybrid of rock lizards of genus Darevskia // 14th European Congress of Herpetology, Porto, Portugal, 19- September, 2007: 306.

20. ., ., . Emys orbicularis // , 2008, 1-2 (60):

183-187.

21. Arakelyan M. Parthenogenesis in Caucasian rock lizard of the genus Darevskia from Armenia. Evolution of parthenogenesis in view of host- parasite relatives // In: Corti C. (ed.) Herpetologia Sardiniae. Soc. Herp. Ital. Edizioni Belvedere, Latina, 2008, 8: 25-28.

22. Arakelyan M., Danielyan F. Parthenogenesis in Caucasian rock lizard of genus Darevskia // In: Corti C. (ed.) Herpetologia Sardiniae. Soc. Herp. Ital. Edizioni Belvedere, Latina, 2008, 8: 29-32.

23. Arakelyan M., Parham J. The geographic distribution of turtles in Armenia and Nagorno-Karabakh Republic (Artsakh) // Chelonian Conservation and Biology, 2008, 7(1): 7077.

24. Arakelyan M., Parham J., Turkozan O., Danielyan F. Sympatrisches vorkommen zweier formen von Testudo graeca in Armenien und der Republik Nagorno Karabakh // Marginata, 2008: 26-30.

25. Danielyan F., Arakelyan M., Stepanyan I. The progress of microevolution in hybrids of rock lizards of genus Darevskia // Biological Journal of Armenia, 2008, 1-2 (60): 147-156.

26. Danielyan F., Arakelyan M., Stepanyan I. Hybrids of Darevskia valentini, D.

armeniaca and D. unisexualis from a sympatric population in Armenia // Amphibia-Reptilia, 2008, 29: 487-504.

27. Arakelyan M., Stepanyan I. The size of red cells of hybrids and parental species of lizards of genus Darevskia // 6th Symposium on the Lacertids of the Mediterranean Basin, Mythimna Lesvos, Greece, 23-27 June, 2008: 15.

28. Arakelyan M., Danielyan F., Hezaveh N. Zoogeographical and taxonomic analysis of Emys orbicularis from Arax River Valley // 15th National and 3rd International conference of Biology, University of Tehran, Iran, 19-21 Aug, 2008: 132.

29. Arakelyan M., Danielyan F., Turkozan O., Parham J. Testudo graeca armeniaca from Armenia and NKR // 15th European Congress of Herpetology, Kuadas Aydin, Turkey, 28 September-2 October, 2009:14.

30. Danielyan F., Arakelyan M. Conservation measures for endangered endemic species of lizards in Armenia // 15th European Congress of Herpetology, Kuadas Aydin, Turkey, 28 September-2 October, 2009: 127.

31. Arakelyan M., Soghoyan A., Ilgaz., Kumluta Y., Durmu S., Tayhan Y.

Morphological and skeletochronological studies on four parthenogenetic species of genus Darevskia // 7th International Symposium on the Lacertids of the Mediterranean basin, Palma, Balearic Islands, Spain, 06-09 September, 2010: 62.



32. Arakelyan M., Danielyan F., Corti C., Sindaco R., Leviton A. The herpetofauna of Armenia and Nagorno Karabakh // In: Di Tizio L., Di Cerbo A.R., Di Francesco N., Cameli A. (eds). Atti VIII Convegno Nazionale Societas Herpetologica Italica, Ianieri Edizioni, Pescara, Chieti, 22-26 September, 2010: 30-32.

33. Freitas S., Perera A., Harris D., Arakelyan M., Danielyan F., Corti C., Carretero M. Phylogeography of the Darevskia raddei species complex based on mtDNA // 7th International Symposium on the Lacertids of the Mediterranean basin, Palma, Balearic Islands, Spain, 06-09 September, 2010: 54.

34. ., . // a. , , 26- , 2011: 42-44.

35. Arakelyan M., Danielyan F., Corti C., Sindaco R., Leviton A. Herpetofauna of Armenia and Nagorno-Karabakh // Salt Lake City SSAR, USA, 2011: 154.

36. Arakelyan M., Danielyan F. Syntopic populations of Darevskia biparental species in Armenia and Nagorno-Karabakh // SEH European Congress of Herpetology and DGHT Deutscher Herpetologentag, Luxembourg and Trier, 25-29 September, 2011: 25-26.

37. Carretero M., Sillero N., Corti C., Jorge F., Freitas S., Arakelyan M. Unkenreflex in Rana macrocnemis from Armenia // Herpetology Notes, 2011, 4: 067-069.

38. Maia J., Ftima J., Perera A., Carretero M., Arakelyan M., Harri J. Molecular assessment of hemoparasites from European and North African Reptiles // 2nd Mediterranean Congress of Herpetology, Marrakech, Morocco, 23-27 May, 2011:

66.

39. Mashkaryan V., Arakelyan M., Harris D., Carretero M. Genetic diversity of Testudo graeca in Armenia and Nagorno-Karabakh Republic // SEH European Congress of Herpetology and DGHT Deutscher Herpetologentag, Luxembourg and Trier, 25-29 September, 2011: 48.

40. Arakelyan M. Conservation status of the last Armenian population of racerunner Eremias arguta transcaucasica // Biological Journal of Armenia 1(64), 2012: 6-9.

41. Arakelyan M. Age and growth of Ophisops elegans from Armenia // Electronic Journal of Natural Sciences, 19 (2), 2012: 63-67.

42. Arakelyan M. Biodiversity of Araks River Valley // 12th International Congress on the Zoogeography and Ecology of Greece and Adjacent Regions, Athens, Greece, 18-22 June, 2012: 17.

Testudo Emys, Darevskia Lacerta,, :

( - Cyt-b ), ( ), (Testudo graeca ibera) (T. g. armeniaca):, ` :

, :

, Emys orbicularis persica E. o.

orbicularis, :

Darevskia,,, Darevskia :, D. raddei D. portschisnkii : D. valentini x D. unisexualis D. raddei x D. rostombekowi :

, (, ), Darevskia, :, D. unisexualis D. uzzelli :, D. raddei nairensis, :, D. bendimahiensis D. raddei vanensis- : D. sapphirina Darevskia :

D. raddei : D. r.

raddei D. r. nairensis :

D. raddei, D. raddei :

,, :

Lacerta, :

:

ARAKELYAN MARINE Microevolutionary processes in sympatric populations of some reptiles from Republic of Armenia and adjacent territories SUMMARY Current work addresses several fundamental questions in the study of micro evolutionary processes of reticulate evolution based on: a) mechanisms of sympatric speciation among closely related species of lizards and turtles from Armenia and adjacent territories, b) identification the role of hybridization in evolution, and c) estimation of evolutional potential of unisexual species, which are a result of hybridization events.

Our data confirmed the role of sympatric speciation in the populations of four groups of reptiles, e.g. turtles of genus Testudo and Emys, lizards of genus Darevskia and Lacerta from Armenia and adjacent territories. The main factors involved in successful development of sympatic speciation were established and two out of three potential pathways of hybrid speciation have been confirmed.

Morphometric (53 characters of shell) and molecular data (Cyt-b marker of mtDNA) suggests the existence of introgressive hybridization of Mediterranean tortoise Testudo graeca in Araks River Valley (south Armenia and Nagorno-Karabakh Republic (NKR)), where gene flow from T. g. ibera to T. g. armeniaca genotype occurs. The endemic morphotypes of Armenian subspecies T. g. armeniaca in Araks River Valley demonstrate high morphological plasticity coupled with the conditions in their environment. Thus, the highly specialized T. g. armeniaca morphotype with low domed shells were found in sites with soft soil favorable for burrowing in Ararat Valley, whereas on territory of NKR T. g. armeniaca were found alongside individuals that are apparently intermediate with T. g. ibera.

The area of the pond turtle, Emys orbicularis persica was confirmed to reach as far as Araks River Valley on the territory of Armenia, NKR and Iran, where the presence of the sympatric zone with between E. o. persica and E. o. orbicularis in NKR is highly probable.

The rock lizards of genus Darevskia are well known as example of hybrid speciation by way of polyploidy. Current study successfully determined the occurrence of fertile triploid hybrids and a tetraploid individual in zone of syntopy of parthenogenetic and bisexual species of genus Darevskia, capable of hybrid speciation by polyploidy, thus confirming the development of reticulate evolution in this genus. It was shown that triploid hybrids have heterotic vigor, while tetraploid has none.

Comparison of different combinations of species of Darevskia in different syntopic zones have revealed that intensive microevolution takes place in this sympatric populations of biparental species D. raddei and D. portschinskii, producing offspring with intermediate scalation and coloration characters. The most successful pairs on second phase of reticulate evolution are biparental D. valentini and parthenogenetic D.

unisexualis as well as biparental D. raddei and parthenogenetic D. rostombekowi, producing fertile hybrids.

Our morphological, skeletochronological and genetic studies have shown that each parthenogenetic species of Darevskia arise from hybridization event of different pairs of maternal and paternal species from different populations. Thus, in spite of similarity of morphology of D. unisexualis and D. uzzelli, whose mother population was D. r. nairensis from slopes of Mt. Aragats, the paternal species are differ. The maternal population of parthenogenetic species D. bendimahiensis is D. r. vanensis from vicinity of Van Lake. Contrary to other studied Darevskia species, D. sapphirina has narrowest periosteal bone which is special mutation of origin and also differs by morphology from other parthenogenetic species.

The D. raddei complex was shown to be of high morphological and genetic variability and its monophylic origin was confirmed. The dynamics of the reproductive development of two closely related forms raddei and nairensis did not specify any significant changes in the duration of reproductive developments, suggesting that raddei is paraphyletic to nairensis. The phylogenetic analyses of D. raddei complex proved an evidence of succession of this species from Araks valley towards north and west.

Comparison of infected lizards from sympatric populations has revealed that parthenogenetic species have a lower rate of infection by blood parasites, than its biparental relative species. The high level of heterozygosity (a consequence of their hybrid origin), as well as generation of new clones through advantageous gene mutation being the alternative mechanism to genetic recombination of biparental species assists in host-parasite co-evolution. Thus, parthenogenetic species Darevskia have high evolutional potential because of higher resistance to parasites and the higher potential in exploitations of new territories.

Our data suggests that post-copulative, rather then pre-copulative reproductive isolation is responsible for impaired success of hybridization of lizards of genus Lacerta. The lower survival rate of the hybrids of first generations and the absence of the hybrids of second generation as a result of the incompatibility of genetic material is a decisive factor in sympatric speciation of lizards.

The current study support new grounds in explanation of the mechanisms of rapid speciation during reticulate evolution. Our findings allow the development of net picture of phylogenetic connections, which include hybridization between closely related forms as one of the underlying evolutionary mechanisms.



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