The effects of contact zone on phylogenetic characters in Persian Wheatear

Atefeh Chamani1 Department of Environment

Faculty of Agricultural and Natural Resources, Islamic Azad University, Khorasgan Branch

Isfahan, IRAN e-mail: a.chamani@khuisf.ac.ir

Mohammad Kaboli2 Department of Fisheries and Environmental Sciences

University of Tehran Tehran, IRAN

Mansour Aliabadian3

Department of Biology Ferdowsi University of Mashhad

Mashhad,Iran

Mahamoud Karami4 Department of Fisheries and Environmental Sciences

University of Tehran Tehran, IRAN

Abstract- The breeding ranges of Kurdish Wheatear (Oenanthe xanthoprymna ) and Persian Wheatear (Oenanthe chrysopygia ) in west and North West of Iran overlap and intermediate color variants can be found. It is possible that this supposed hybridization differs the characters of O.chrysopygia between contact zone and other distribution ranges in Iran. We aimed the survey of this possibility in this study. Field works carried out in spring and summer 2006 and 2007 and we found O.chrysopygia inside and outside of contact zone in west, northwest, east and center of Iran. Then, we used DNA sequences of two mitochondrial genes, NADH subunit 2 (ND2); 975 bp and cytochrome oxidase subunit I gene (cox1); 652 bp from 39 individuals and Bayesian, Maximum Likelihood (ML) also Maximum Parsimony (MP) methods to derive a phylogeny for comparison between O.chrysopygia inside and outside of contact zone. Our results confirmed O.chrysopygia in contact zone, well differentiated from O.chrysopygia in outside of contact zone. Furthermore, we confirmed the close phylogenetic relationships of supposed hybrid with O.chrysopygia despite a plumage coloration pattern close to O. xanthoprymna.

Keywords-Wheatear;O.xanthoprymna;O.chrysopygia; supposed hybrid; hybridization; contact zone

I. INTRODUCTION Genus Oenanthe bears considerable interspecific

morphological (wing shape and plumage coloration) variation [3, 9]. Such variation has led to difficulty in assessing taxonomic position of its species [24]. Hybridization occurs between several species in genus Oenanthe like O.finschii and O.picata [11, 7, 14, 8], also between three populations (capistrata, opistholeuca, picata) treated as races of Oenanthe picata [11, 8]. Hybridization between Kurdish Wheatear (O.xanthoprymna) and Persian Wheatear (O.chrysopygia) also is subject of few hypothesis and taxonomic obscurity [3, 16, 10, 19, 8] and are usually recognized by the majority of taxonomists. Persian Wheatear

has grayish under parts and red tail in both male and female coloration. In Kurdish Wheatear the male has brown-black throat and under the wings, red tail and white basal in two-thirds of rectrices, while the female is like Persian Wheatear. Persian Wheatear breeds in the inner Zagros Mountains in all over the Iran [3, 18,10] and is a montane species and inhabits rocky slopes covered with scree, above 2000 m elevation [16]. Breeding range of the Kurdish Wheatear extends from the extreme south-eastern parts of Turkey further south-east, into the Zagros Mountains and probably in extreme north-east Iraq [4].There is probably a fairly narrow zone in western IRAN, where the breeding ranges of these two taxa overlap, interbreeding takes place and birds with intermediate color variants (has been described as O.cummingi ) can be found [3]. The characters found in this intermediate population, are present in few first year birds of Kurdish wheatear [4, 13]. Moreover these variants and Kurdish wheatear are reported at the same localities [21], apparently additional evidence that has been exaggerated about the incidence of hybrids or intermediates [1]. But McCarthy [8] suggested this hybrid population has the black throat of O.xanthoprymna and the red cornered tail of O.chrysopygia and believe that due to hybridization, these birds are sometimes lumped. In this paper, we report on phylogenetic relationships between O.chrysopygia inside and outside of contact zone to identify the effects of contact zone on phylogenetic characters of O.chrysopygia. Furthermore we survey the phylogenetic relationships of O.chrysopygia and supposed hybrid.

II. MATERIAL AND METHOD

A. Sampling and Field works Field works carried out in spring and summer 2006 and

2007 in south, south west, west, northwest, center and north east of Iran and we looked for all suitable habitats for our

V2-287

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

B

taxa inside adistribution ra

We took fduring 2 sa(Kermanshah took O.chrysoKhorasan, 3 scenter (IsfahaOenanthe alb

B. Sequence Phylogene

different optimand (2) Maxiclade supportparametric bPartitioned Bwith MrBayeperformed unaddition repl

Figure 1. Distr

and outside oange in [10] (Ffifteen O.chryampling yeaProvince, Am

opygia outsidespecimens), wan, 10 specimoniger as outg

analysis etic analysesmality criteriaimum Likeliht for the ML

bootstrapping Bayesian Markes 3.1.1 [17, 1nder the heurisicates with t

ribution ranges of

of contact zoFig. 1). ysopygia and ars in contamrolah Regioe of contact zowest (Kurdistamens).We addegroup from [22

s were conda: (1) Maximuhood (ML) usL analyses wa

(100 replicakov Chain M12, 20]. Parsimstic search optthe branch-sw

f O.xanthoprymnabla

one based on

10 supposed hact zone inon). Furthermoone in east (Noan, 3 specimened measurem2].

ducted using um Parsimonysing PAUP*bas assessed bates) [15] al

Monte Carlo mmony analysetion with 10 r

wapping optio

a, O.chrysopygiaack triangles and

n their

hybrids n west ore we orthern ns) and

ments of

three y (MP) 10 [5], y non-lso (3) method, es were random on tree

bino10esLifowiweprge

97phanTh

a and contact zonesampling places

section and reodes was estim000 replicatesstimated usingikelihood Rati

our MCMC cith trees sampere discarderobabilities wenerations.

IThe final ed

75 bp for ND2hylogenetic trenalyses of thehere are three

e (Distribution rashown with black

econnection (Tmated using ths. Likelihood g Modeltest vio Test criterihains were ru

pled every 10 gd (“burn-in”

were estimate

III. RESULTS

dited data set2. There is a gees resulting fe combined dmajor clades i

ange of supposed k circles.

TBR) implemhe non-parame

models andv.3.04 [6] andion. For the Bun for one mgenerations. T” period) and for the re

S AND DISCUS

t included 652general congrufrom the Baye

data set (CO1 in these trees

hybrid); Field wo

mented. Supporetric bootstrapd parameters d the HierarcBayesian anal

million generaThe first 5000 nd the postemaining sam

SION 2 bp for CO1uence betweeesian, ML and

and ND2 ge(Fig. 2).

ork ranges shown

rt for with were

chical lyses, ations trees

terior mpled

1 and en the d MP enes).

n with

V2-288

Figure 2. ML, Mthe Bayesian an

The relatiosimilar in thediffer often iprobabilities iincludes the sCenter of Iranand ML suppgenetically diclade include

MP and Posteriornalysis are indicat

onships withine individual gnvolve short in at least onespecimens of n (outside of cort (100%) anifferentiated frs O.chrysopyg

r probability valuted at the >99% (

n each of the gene trees, an

nodes that ree of the gene tO.chrysopygi

contact zone) nd Posterior prom the secongia in contact

ues from the Maxi**) >95% (*) sig

three clades and the nodes teceive low potrees. The firsia in East, Wethat with strorobability of 0

nd clade. The t zone and su

imum Likelihoodgnificance levels.

discussed in the

are also that do osterior st clade est and

ong ML 0.85, is second

upposed

hyhi(1diprtaxbo

d, Maximum ParsML and MP Boo

e text.

ybrid that welgh bootstrap .00). There afferentiated c

robability) thaxa with immaootstrap (100%

simony and Bayesotstrap values Giv

ll differentiatesupport (10

are only threclade (100% at includes twoature specime

%) and Posteri

sian analysis. Posven only if >90%

ed from the t0%) and Po

ee specimens bootstrap an

o supposed hyens of O.chryior probability

sterior Probability. Clades 1, 2 and

third clade, wsterior probabin the third

nd 1.00 Postybrid that are ysopygia with y (1.00).

y from 3 are

with a bility well

terior sister high

2011 2nd International Conference on Environmental Science and Technology (ICEST 2011)

V2-289

IV. CONCLUSION We confirm, with a high posterior probability and

bootstrap support that O. chrysopygia in contact zone, well differentiated from O.chrysopygia in outside of contact zone. Then, we suggest that overlapping between distribution ranges of O. chrysopygia and O. xanthoprymna in western Iran, makes differences the phylogenetic characters of O.chrysopygia in contact zone with O.chrysopygia in West, East and Center of Iran that located in outside of contact zone. These differences can be the impacts of hybridization incidence in contact zone. Furthermore, the Supposed hybrids have close phylogenetic relationships with O.chrysopygia, despite a plumage pattern close to O. xanthoprymna.

ACKNOWLEDGMENT We thank Eric.Pasquet in Muséum National d’Histoire

Naturelle, Département Systématique et Evolution of Paris to help us in molecular analysis and also gratefully acknowledge the Iran Department of Environment for providing scientific collecting permits.

REFERENCES [1] Association of European Records and Rarities Committees (AERC).

AERC Taxonomic Recommendations, 2003, Available from: http://www.aerc.be/aerc_tac.htm .

[2] B.T. Winnepenninckx, R. Backeljau and De. Wacheter, “Extraction of high molecular weight DNA for mollusk”. TIG. 9(12), 1993, p:407.

[3] C. Vaurie, “Notes on the bird genus Oenanthe in persica, Afghanistan and India”. No.1425, 1949.

[4] C.S. Roselaar, “Taxonomy - Morphology - Distribution songbird of Turkey: an atlas of biodiversity of Turkish passerine birds”, Pica press, London, 1995.

[5] D.L. Swofford, PAUP*. “Phylogenetic Analysis Using Parsimony (*and other methods)”, Version 4b10. Sinauer Associates, Sunderland, Massachusets, 2002.

[6] D. Posada and K.A. Crandall, “Modeltest: Testing the model of DNA substitution”. Bioinformatics. 14, 1998, pp:817-818.

[7] E.C. Dickinson, The Howard and Moore Complete Checklist of the Birds of the World”. London, Christopher Helm, 2003.

[8] E.M. McCarthy, “Handbook of Avian Hybrids of the World”. New York, NY, Oxford University Press, 583 pp., ISBN, 0-19-518323-1, 2006.

[9] E. Mayr and E. Stresemann, “Polymorphism in the Chat genus Oenanthe (Aves) ”.Evolution. 4, 1955, pp: 291-300.

[10] E.N. Panov, “Wheatears of Palaearctic: ecology, behavior and evolution of the genus Oenanthe”, Pensoft, Sofia-Moscow, 2005.

[11] E.N. Panov, V.I. Grabovsky, S.V. Ljubustchenko, “Divergence and hybridpolymorphism in the complex “Eastern Pied Wheatears”, Oenanthe picata”. Zool Zh. 72, 1993, pp: 80–96.

[12] F. Ronquist, J.P. Huelsenbeck, “MrBayes 3: Bayesian phylogenetic inference under mixed models”. Bioinformatics. 19, 2003, pp:1572–1574.

[13] G.L. Bates, On O.xanthoprymna & O.chrysopygia. Ibis. (13)5,1935, pp:198-201

[14] J. Del Hoyo, A. Elliott and D. A. Christie, “Handbook of the Birds of the World” V.10. Spain, Lynx Edicions, 2005.

[15] J. Felsenstein, “Confidence limits on phylogenies: an approach using bootstrap”. Evolution. 39, 1985, pp:783–791.

[16] J. Haffer, “Secondary contact zones of birds in northern Iran,” Bonn. Zool. Monogr. 10, 1977, pp:1-41.

[17] J. P. Huelsenbeck and F. Ronquist, “MrBayes: Bayesian inference of phylogenetic trees”. Bioinformatics. 17, 2001, pp:754–755.

[18] L. Cornwallis, “The comparative ecology of eleven species of wheatear (genus Oenanthe) in S. W. Iran,” Ph.D thesis, Oxford University.

[19] M. Aliabadian, M.Kaboli, R.Prodon, V. Nijman and M.Vences, “Phylogeny of Palaearctic wheatears (genus Oenanthe) – congruence between morphometric and molecular data,” Mol. Phyl. Evol. 42(3), 2007, pp: 665-675.

[20] M.C. Brandley, A. Schmitz, TW. Reeder, 2005. “Partitioned Bayesian analyses, partition choice, and the phylogenetic relationships of scincid lizards”. Syst. Biol. 54, 373-390.

[21] M. Harms, “Oenanthe xanthoprymna (Hemprich & Ehrenberg),” J.Orn. 73, 1925, pp:390-394.

[22] M. Kaboli, M. Aliabadian, K. S. Guillaumet, K. Roselaar and R. Prodon, “Ecomorphology of the wheatears (genus Oenanthe),” Ibis. 149, 2007, pp:792-805.

[23] P.D.N. Herbert, M.A. Stoeckle, T.S. Zemlak and C.M. Francis, “ Identification of birdsthrough DNA barcodes,” PLOS Biol. vol 2, 2004, pp.1657-1663.

[24] R.K. Outlaw, G. Voelker and R.C.K. Bowie, “Shall we chat? Evolutionary relationships in the genus Cercomela (Muscicapidae) and its relation to Oenanthe reveals extensive polyphyly among chats distributed in Africa, India and the Palearctic”. Mol. Phyl. Evol. 55(1), 2010, pp: 284-92.

[25] R.M. Zink, S. Drovetski and S. Rohwer, “Phylogeographic patterns in the Great Spotted Woodpecker (Dendrocopos major) across Eurasia”. J.Avian Biol. 35, 2002, pp: 175–178.

2011 2nd International Conference on Environmental Science and Technology (ICEST 2011)

V2-290