Research Article Phylogeography of crossbills, bullfinches, grosbeaks, and rosefinches A.Arnaiz-Villenaa,*, J. Guilléna,V. Ruiz-del-Vallea, E. Lowy, J. Zamoraa, P.Varelaa, D. Stefanib and L.M.Allendea aDepartment of Immunology and Molecular Biology, Hospital 12 de Octubre, Universidad Complutense, 28041 Madrid (Spain), Fax +34 91 390 83 99, e-mail: aarnaiz@eucmax.sim.ucm.es bviale Colombo, 67, Cagliari (Sardinia) Received 26 March 2001; received after revision 7 May 2001; accepted 6 June 2001 Abstract. Mitochondrial cytochrome b (cyt b) from 24 Carduelini   species   including   crossbills,   bullfinches, grosbeaks, rosefinches, and other related, but not conclu- sively   classified   species,   was   sequenced.   These   se- quences  were  also  compared  with  all  the  available  se- quences from the genera Carduelis, Serinus, and Passer. Phylogenetic analyses consistently gave the same groups of  finches  and  the  calculated  divergence  times  suggest that speciation of the studied species occurred between 14 and 3 million years ago (Miocene-Pliocene), appear- ing  before  the  Passer,  Carduelis,  and  Serinus  genera. Pleistocene glaciations may have been important in sub- speciation.  Crossbills  are  integrated  within  the  genus Carduelis, together with redpolls; the common crossbill some other related genera comprising one or two species [1].  DNA  hybridization  techniques  have  placed  Car- duelini  finches  closely  to  Hawaiian  honeycreepers  [1]. However, this technique presents many difficulties in in- terpretation and the Carduelini have also been related to ploceids and estrilids by osteological parameters [2] and to chaffinches, bramblings, and honeycreepers using my- ological data [3]. In the present work, we aimed to study the phylogenetic relationships among Carduelini (particularly crossbills, bullfinches,   grosbeaks,   and   rosefinches)   at   the   mi- croevolutionary    level    by    using    mitochondrial    cy- CMLS, Cell. Mol. Life Sci. 58 (2001) 001–08 1420-682X/01/010001-08 $ 1.50 + 0.20/0 © Birkhäuser Verlag, Basel, 2001 CMLS Cellular and Molecular Life Sciences CMLS 1163 shows  subspeciation  with  Loxia  japonica  in  the  Pleis- tocene epoch. Pinicola enucleator  groups together with bullfinches  and  is  probably  the  ancestor  of  the  group. Hawfinch is only distantly related to the studied groups, and might either represent an isolated genus or be related to  the  New  World  genus  Hesperiphona.  The  grosbeak genera Eophona and Mycerobas are clearly sister groups, and species belonging to the former might have given rise to  Mycerobas  species.  The  isolated  (in  classification) Uragus sibericus and Haematospiza sipahi  are included within the genus Carpodacus (rosefinches); Carpodacus nipalensis is outside the genus Carpodacus in the molec- ular analyses and might be an isolated species or related to the genus Montifringilla. Key words. Crossbill; bullfinch; grosbeaks, rosefinch; goldfinch; redpoll; Carduelini; canary; sparrow; greenfinch; siskin; linnet. The  Order  Passeriformes  includes  the  Passerine  birds, and contains several families and subfamilies represent- ing about one-half of extant birds [1]. The classification, phylogeny, and relationships of the so-called Carduelini finches remain unresolved. The tribe Carduelini includes goldfinches  (genus  Carduelis),  crossbills  (Loxia),  ca- naries  (Serinus), rosefinches (Carpodacus), bullfinches (Pyrrhula)  and  grosbeaks  (Eophona,  Mycerobas)  and Corresponding author. A. Arnaiz-Villena and J. Guillén contributed equally to this work.