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) 00108
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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.