2011 2nd International Conference on Environmental Science and Technology IPCBEE vol.6 (2011) © (2011) IACSIT Press, Singapore

The effects of contact zone on phylogenetic characters in Persian

Atefeh Chamani1 Mansour Aliabadian3 Department of Environment Department of Biology Faculty of Agricultural and Natural Resources, Ferdowsi University of Mashhad Islamic Azad University, Khorasgan Branch Mashhad, Isfahan, IRAN e-mail: [email protected]

Mohammad Kaboli2 Mahamoud Karami4 Department of Fisheries and Environmental Sciences Department of Fisheries and Environmental Sciences University of Tehran University of Tehran Tehran, IRAN Tehran, IRAN

Abstract- The breeding ranges of Kurdish Wheatear (Oenanthe has grayish under parts and red tail in both male and female xanthoprymna ) and Persian Wheatear (Oenanthe chrysopygia ) coloration. In Kurdish Wheatear the male has brown-black in west and North West of Iran overlap and intermediate color throat and under the wings, red tail and white basal in two- variants can be found. It is possible that this supposed thirds of rectrices, while the female is like Persian Wheatear. hybridization differs the characters of O.chrysopygia between Persian Wheatear breeds in the inner Zagros Mountains in all contact zone and other distribution ranges in Iran. We aimed over the Iran [3, 18,10] and is a montane species and inhabits the survey of this possibility in this study. Field works carried rocky slopes covered with scree, above 2000 m elevation out in spring and summer 2006 and 2007 and we found [16]. Breeding range of the Kurdish Wheatear extends from O.chrysopygia inside and outside of contact zone in west, the extreme south-eastern parts of further south-east, northwest, east and center of Iran. Then, we used DNA into the Zagros Mountains and probably in extreme north- sequences of two mitochondrial genes, NADH subunit 2 (ND2); 975 bp and cytochrome oxidase subunit I gene (cox1); 652 bp east [4].There is probably a fairly narrow zone in from 39 individuals and Bayesian, Maximum Likelihood (ML) western IRAN, where the breeding ranges of these two taxa also Maximum Parsimony (MP) methods to derive a phylogeny overlap, interbreeding takes place and with for comparison between O.chrysopygia inside and outside of intermediate color variants (has been described as contact zone. Our results confirmed O.chrysopygia in contact O.cummingi ) can be found [3]. The characters found in this zone, well differentiated from O.chrysopygia in outside of intermediate population, are present in few first year birds of contact zone. Furthermore, we confirmed the close Kurdish wheatear [4, 13]. Moreover these variants and phylogenetic relationships of supposed hybrid with Kurdish wheatear are reported at the same localities [21], O.chrysopygia despite a plumage coloration pattern close to O. apparently additional evidence that has been exaggerated xanthoprymna. about the incidence of hybrids or intermediates [1]. But McCarthy [8] suggested this hybrid population has the black Keywords-Wheatear;O.xanthoprymna;O.chrysopygia; throat of O.xanthoprymna and the red cornered tail of supposed hybrid; hybridization; contact zone O.chrysopygia and believe that due to hybridization, these birds are sometimes lumped. In this paper, we report on I. INTRODUCTION phylogenetic relationships between O.chrysopygia inside and Genus Oenanthe bears considerable interspecific outside of contact zone to identify the effects of contact zone morphological (wing shape and plumage coloration) on phylogenetic characters of O.chrysopygia. Furthermore variation [3, 9]. Such variation has led to difficulty in we survey the phylogenetic relationships of O.chrysopygia assessing taxonomic position of its species [24]. and supposed hybrid. Hybridization occurs between several species in genus Oenanthe like O.finschii and O.picata [11, 7, 14, 8], also II. MATERIAL AND METHOD between three populations (capistrata, opistholeuca, picata) A. Sampling and Field works treated as races of Oenanthe picata [11, 8]. Hybridization between Kurdish Wheatear (O.xanthoprymna) and Persian Field works carried out in spring and summer 2006 and Wheatear (O.chrysopygia) also is subject of few hypothesis 2007 in south, south west, west, northwest, center and north and taxonomic obscurity [3, 16, 10, 19, 8] and are usually east of Iran and we looked for all suitable habitats for our recognized by the majority of taxonomists. Persian Wheatear

V2-287 taxa inside and outside of contact zone based on their bisection and reconnection (TBR) implemented. Support for distribution range in [10] (Fig. 1). nodes was estimated using the non-parametric bootstrap with We took fifteen O.chrysopygia and 10 supposed hybrids 1000 replicates. Likelihood models and parameters were during 2 sampling years in contact zone in west estimated using Modeltest v.3.04 [6] and the Hierarchical (Kermanshah Province, Amrolah Region). Furthermore we Likelihood Ratio Test criterion. For the Bayesian analyses, took O.chrysopygia outside of contact zone in east (Northern four MCMC chains were run for one million generations Khorasan, 3 specimens), west (Kurdistan, 3 specimens) and with trees sampled every 10 generations. The first 5000 trees center (Isfahan, 10 specimens).We added measurements of were discarded (“burn-in” period) and the posterior Oenanthe alboniger as outgroup from [22]. probabilities were estimated for the remaining sampled generations. B. Sequence analysis Phylogenetic analyses were conducted using three different optimality criteria: (1) Maximum Parsimony (MP) III. RESULTS AND DISCUSSION and (2) Maximum Likelihood (ML) using PAUP*b10 [5], The final edited data set included 652 bp for CO1 and clade support for the ML analyses was assessed by non- 975 bp for ND2. There is a general congruuence between the parametric bootstrapping (100 replicates) [15] also (3) phylogenetic trees resulting from the Bayesian, ML and MP Partitioned Bayesian Markov Chain Monte Carlo method, analyses of the combined data set (CO1 and ND2 genes). with MrBayes 3.1.1 [17, 12, 20]. Parsimony analyses were There are three major clades iin these trees (Fig. 2). performed under the heuristic search option with 10 random addition replicates with the branch-swapping option tree

Figure 1. Distribution ranges of O.xanthoprymna, O.chrysopygia and contact zone (Distribution range of supposed hybrid); Field work ranges shown with black triangles and sampling places shown with black circles.

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Figure 2. ML, MP and Posterior probability values from the Maximum Likelihood, Maximum Parsimony and Bayesian analysis. Posterior Probability from the Bayesian analysis are indicated at the >99% (**) >95% (*) significance levels. ML and MP Bootstrap values Given only if >90%. Clades 1, 2 and 3 are discussed in the text.

The relationships within each of the three clades are also hybrid that well differentiateed from the third clade, with a similar in the individual gene trees, and the nodes that do high bootstrap support (100%) and Posterior probability differ often involve short nodes that receive low posterior (1.00). There are only three specimens in the third well probabilities in at least one of the gene trees. The first clade differentiated clade (100% bootstrap and 1.00 Posterior includes the specimens of O.chrysopygia in East, West and probability) that includes two supposed hybrid that are sister Center of Iran (outside of contact zone) that with strong ML taxa with immature specimens of O.chrysopygia with high and ML support (100%) and Posterior probability of 0.85, is boootstrap (100%) and Posterior probability (1.00). genetically differentiated from the second clade. The second clade includes O.chrysopygia in contact zone and supposed

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IV. CONCLUSION [9] E. Mayr and E. Stresemann, “Polymorphism in the Chat genus Oenanthe (Aves) ”.Evolution. 4, 1955, pp: 291-300. We confirm, with a high posterior probability and [10] E.N. Panov, “ of Palaearctic: ecology, behavior and bootstrap support that O. chrysopygia in contact zone, well evolution of the genus Oenanthe”, Pensoft, Sofia-Moscow, 2005. differentiated from O.chrysopygia in outside of contact zone. [11] E.N. Panov, V.I. Grabovsky, S.V. Ljubustchenko, “Divergence and Then, we suggest that overlapping between distribution hybridpolymorphism in the complex “Eastern Pied Wheatears”, ranges of O. chrysopygia and O. xanthoprymna in western Oenanthe picata”. Zool Zh. 72, 1993, pp: 80–96. Iran, makes differences the phylogenetic characters of [12] F. Ronquist, J.P. Huelsenbeck, “MrBayes 3: Bayesian phylogenetic O.chrysopygia in contact zone with O.chrysopygia in West, inference under mixed models”. Bioinformatics. 19, 2003, pp:1572– East and Center of Iran that located in outside of contact 1574. zone. These differences can be the impacts of hybridization [13] G.L. Bates, On O.xanthoprymna & O.chrysopygia. Ibis. (13)5,1935, pp:198-201 incidence in contact zone. Furthermore, the Supposed [14] J. Del Hoyo, A. Elliott and D. A. Christie, “Handbook of the Birds of hybrids have close phylogenetic relationships with the World” V.10. Spain, Lynx Edicions, 2005. O.chrysopygia, despite a plumage pattern close to O. [15] J. Felsenstein, “Confidence limits on phylogenies: an approach using xanthoprymna. bootstrap”. Evolution. 39, 1985, pp:783–791. [16] J. Haffer, “Secondary contact zones of birds in northern Iran,” Bonn. ACKNOWLEDGMENT Zool. Monogr. 10, 1977, pp:1-41. We thank Eric.Pasquet in Muséum National d’Histoire [17] J. P. Huelsenbeck and F. Ronquist, “MrBayes: Bayesian inference of Naturelle, Département Systématique et Evolution of Paris to phylogenetic trees”. Bioinformatics. 17, 2001, pp:754–755. help us in molecular analysis and also gratefully [18] L. Cornwallis, “The comparative ecology of eleven species of acknowledge the Iran Department of Environment for wheatear (genus Oenanthe) in S. W. Iran,” Ph.D thesis, Oxford University. providing scientific collecting permits. [19] M. Aliabadian, M.Kaboli, R.Prodon, V. Nijman and M.Vences, “Phylogeny of Palaearctic wheatears (genus Oenanthe) – congruence EFERENCES R between morphometric and molecular data,” Mol. Phyl. Evol. 42(3), [1] Association of European Records and Rarities Committees (AERC). 2007, pp: 665-675. AERC Taxonomic Recommendations, 2003, Available from: [20] M.C. Brandley, A. Schmitz, TW. Reeder, 2005. “Partitioned Bayesian http://www.aerc.be/aerc_tac.htm . analyses, partition choice, and the phylogenetic relationships of [2] B.T. Winnepenninckx, R. Backeljau and De. Wacheter, “Extraction scincid lizards”. Syst. Biol. 54, 373-390. of high molecular weight DNA for mollusk”. TIG. 9(12), 1993, [21] M. Harms, “Oenanthe xanthoprymna (Hemprich & Ehrenberg),” p:407. J.Orn. 73, 1925, pp:390-394. [3] C. Vaurie, “Notes on the genus Oenanthe in persica, Afghanistan [22] M. Kaboli, M. Aliabadian, K. S. Guillaumet, K. Roselaar and R. and ”. No.1425, 1949. Prodon, “Ecomorphology of the wheatears (genus Oenanthe),” Ibis. [4] C.S. Roselaar, “ - Morphology - Distribution songbird of 149, 2007, pp:792-805. Turkey: an atlas of biodiversity of Turkish birds”, Pica [23] P.D.N. Herbert, M.A. Stoeckle, T.S. Zemlak and C.M. Francis, press, London, 1995. “ Identification of birdsthrough DNA barcodes,” PLOS Biol. vol 2, [5] D.L. Swofford, PAUP*. “Phylogenetic Analysis Using Parsimony 2004, pp.1657-1663. (*and other methods)”, Version 4b10. Sinauer Associates, Sunderland, [24] R.K. Outlaw, G. Voelker and R.C.K. Bowie, “Shall we chat? Massachusets, 2002. Evolutionary relationships in the genus Cercomela (Muscicapidae) [6] D. Posada and K.A. Crandall, “Modeltest: Testing the model of DNA and its relation to Oenanthe reveals extensive polyphyly among chats substitution”. Bioinformatics. 14, 1998, pp:817-818. distributed in Africa, India and the Palearctic”. Mol. Phyl. Evol. 55(1), [7] E.C. Dickinson, The Howard and Moore Complete Checklist of the 2010, pp: 284-92. Birds of the World”. London, Christopher Helm, 2003. [25] R.M. Zink, S. Drovetski and S. Rohwer, “Phylogeographic patterns in [8] E.M. McCarthy, “Handbook of Avian Hybrids of the World”. New the Great Spotted Woodpecker (Dendrocopos major) across Eurasia”. York, NY, Oxford University Press, 583 pp., ISBN, 0-19-518323-1, J.Avian Biol. 35, 2002, pp: 175–178. 2006.

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