A multilocus phylogeny of New World jay genera

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Abstract

We studied phylogenetic relationships of the New World Jays (NWJs) based on DNA sequences from three mitochondrial and two nuclear loci. Sampling included at least two individuals from each of the seven NWJ genera and four outgroups of closely related corvids, as well as six of the 16 Cyanocorax species (including two representatives of the previously recognized “Cissilopha”). Phylogenetic analyses were conducted using maximum parsimony, maximum likelihood, and Bayesian analyses for individual genes and a combined dataset. The combined phylogenetic analysis supports the basal position of Cyanolyca to all other NWJs, a (Cyanocorax (Calocitta, Psilorhinus)) clade, and a ((Cyanocitta, Aphelocoma) Gymnorhinus) clade that agrees with a novel morphological synapomorphy uniting Cyanocitta and Aphelocoma. Within Cyanocorax, C. yncas (former “Xanthoura”) is basal to a split among former “Cyssilopha” species and the rest of the Cyanocorax species. To explore implications for the historical biogeography of the NJWs, we used Dispersal–Vicariance Analysis, which indicated that NWJs originated either in Mesoamerica or North America + Mesoamerica, with South American NWJs dispersing three times independently from Mesoamerica.

Introduction

The New World jays (NWJs) are an assemblage of 34 species in seven genera of corvids—Aphelocoma, Cyanocitta, Gymnorhinus, Cyanolyca, Calocitta, Psilorhinus, and Cyanocorax—endemic to the Americas. Monophyly of this assemblage is not in serious doubt, given unique shared morphological characters (Zusi, 1987). NWJs have radiated successfully in tropical, subtropical, and temperate habitats, and show a mix of narrow endemism (e.g., Cyanolyca mirabilis) and broad distributions (e.g., Cyanocitta cristata), including species of particular conservation concern (BirdLife International, 2000). Furthermore, they have served as a model system for numerous analyses of the evolution of avian social systems (e.g., Brown, 1963, Fitzpatrick and Woolfenden, 1985, Edwards and Naeem, 1993, Saunders and Edwards, 2000). As such, understanding the historical underpinnings of current NWJ diversity is critical on a number of fronts.

Although a broad older literature attempted to treat NWJ history and evolution (Amadon, 1944, Hardy, 1961), the first steps towards an understanding of NWJ phylogeny began with a study based on cytochrome b sequences (Espinosa de los Monteros and Cracraft, 1997), which provided a basic framework, but little detail of specific relationships. Later studies (Ericson et al., 2005, Saunders and Edwards, 2000) added detail, and considered additional genes in making the NWJ tree more robust.

However, two of the previous studies (Ericson et al., 2005, Espinosa de los Monteros and Cracraft, 1997) considered only single representatives of each genus, providing little detail useful for understanding the historical biogeography of the clade and no test of monophyly of genera. The more detailed study (Saunders and Edwards, 2000), however, considered but a single gene (the mitochondrial control region) and included only two South American NWJ representatives. As such, considerable room remains for clarifying the details of NWJ phylogeny.

The present paper offers a multigene view of the ‘deep’ nodes in the NWJ phylogeny. We have assembled much improved sampling of key species from each genus, particularly as relates to the older phylogeny and biogeography of the group, as well as sequence data from additional mitochondrial and nuclear loci. This study offers a key understructure to understanding the broader picture of NWJ history and evolution.

Section snippets

Sampling

We included at least two individuals from each of the seven NWJ genera—Aphelocoma, Cyanocitta, Calocitta, Psilorhinus, Cyanocorax (including Cissilopha) and Gymnorhinus—and four individuals representing closely related corvid genera: Dendrocitta, Pica, Perisoreus, and Corvus. Also, in a first attempt to elucidate relationships among Cyanocorax jays, we analyzed six of the 16 currently recognized species of Cyanocorax, including two representatives of the previously recognized “Cissilopha

Sequence attributes

DNA sequence lengths and general characteristics for each gene are summarized in Table 1. As expected, sequence variation was comparable among the mitochondrial genes, and substantially higher than in the nuclear introns, with βfib7 evolving faster than AK5. Pairwise distances among the 18 species of jays and 4 outgroups for ND2, cytb, and βfib7, are summarized in Fig. 1. Among interesting features of divergence is that when plotting uncorrected distances, ND2 and cytb saturate early compared

Phylogenetic analyses

Our combined phylogenetic tree (Fig. 5) depicts intergeneric relationships among NWJs. Relatively weak nodes of the tree are restricted to one clade—that including Aphelocoma, Cyanocitta, and Gymnorhinus (the “ACG” clade). The low support for the monophyly of ACG is a consequence of CR, which supports a topology that differs from those of the βfib7, ND2, and cytb trees. Since CR is physically linked to the other mitochondrial loci, its different topology should not be the result of independent

Acknowledgments

We are grateful to A.G. Navarro-Sigüenza (Museo de Zoologı´a, Universidad Nacional Autónoma de México), S. Hackett and D.E. Willard (Field Museum), S. Birks (Burke Museum of Natural History and Culture), and J. Craraft and P. Sweet (American Museum of Natural History), and M.B. Robbins (University of Kansas Natural History Museum) for providing tissue samples under their care. A.G. Navarro-Sigüenza and B. Hernández-Baños made possible sequencing of Cyanocorax dickeyi in Mexico. Two anonymous

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