A phylogeny of extant penguins (Aves: Sphenisciformes) combining morphology and mitochondrial sequences

Abstract

The phylogenetic relationships among the penguins have received little attention, despite their well-known anatomy and the conspicuous nature of the group. Previous attempts have included datasets limited to few, mostly osteological characters, and one study was based on integumentary and breeding characters. We developed a morphological matrix comprising 159 morphological characters of osteology (70 characters), myology (15), digestive tract (1), integument (66), and breeding (7 characters), scored in 18 extant forms (all currently recognized species plus one distinct subspecies). A gaviiform was placed at the root, and 11 species of representative procellariiform groups completed the outgroup. A heuristic parsimony analysis under equal weights was performed. We also compiled DNA sequences available in GenBank for the mitochondrial genes 12S rDNA and cytochrome b. We included the two data partitions in a combined analysis under direct optimization. Both analyses recovered the monophyly of Sphenisciformes and all the traditional polytypic genera. Morphological characters performed optimally at the ordinal and generic nodes, also providing resolution and varying degrees of support at supra-and intrageneric nodes. The comparison of molecular and morphological results indicated that the most significant problem in the phylogeny of extant penguins is rooting the ingroup. The mutual interaction of molecular and morphological data decreases the ambiguity regarding the placement of the root, and provides a resolved, relatively well-supported phylogeny of extant penguins. Biogeographical patterns based on breeding ranges and derived from the combined analysis show that the major intercontinental vicariance events detected are consistent with cold marine current patterns of the Southern Hemisphere.

The Basal Penguin (Aves: Sphenisciformes) Perudyptes devriesi and a Phylogenetic Evaluation of the Penguin Fossil Record

“…Some differences between our results and those of Acosta Hospitaleche et al (2007,2008) can be accounted for by the effect of the different preferred rooting for crown penguins when molecular data are included (discussed by Bertelli and Giannini, 2005). However, reexamination of the character evidence suggests that three major differences are driven by character sampling issues and erroneous scorings.…”

“…This conclusion is also supported by morphological differences and the allopatric distribution of the three species (Banks et al, 2006). In past studies (Giannini and Bertelli, 2005;Bertelli and Giannini, 2006;Clarke et al, 2007), Eudyptes chrysocome chrysocome and Eudyptes chrysocome moseleyi were coded as separate operational taxonomic units in recognition of these morphological differences. Here, we treat all three of the newly recognized rockhopper species as separate terminals.…”

“…Earlier versions of this dataset were formulated and utilized by Giannini and Bertelli (2004), Bertelli and Giannini (2005), Bertelli et al (2006), , Clarke et al (2007), and, most recently, by Ksepka (2007). In the present study, we modified Ksepka's (2007) version of this matrix, including 25 morphological characters and 11 fossil taxa added since the most recent published version of the matrix (Clarke et al, 2007).…”

“…These incomplete specimens pose a challenge to phylogenetic analysis (discussed by Fordyce and Jones, 1990) and have hindered the incorporation of biogeographical, temporal, and morphological data from such taxa into the emerging synthesis of penguin evolution. Herein, we undertake an evaluation of the phylogenetic position of all fossil penguin taxa and multiple sets of remains recognized as representing as yet unnamed taxa, with the goal of improving our understanding of the tempo of penguin evolution and contributing to the debate regarding the timing of the radiation of crown Spheniscidae (Bertelli and Giannini, 2005;Baker et al, 2006;Jadwiszczak, 2006b;Clarke et al, 2007).…”

A Cryptic New Species of Thrush (Turdidae:Turdus) from Western Amazonia

“…Recently, the integration of molecular techniques, vocal analysis, and traditional morphology has facilitated, more than ever, discovery of new species (e.g., Krabbe et al 2005, O'Neill et al 2011. In particular, improvements in the methods and cost-effectiveness of molecular analyses, in conjunction with increased higher education opportunities for Neotropical ornithologists, have significantly augmented the contributions of native ornithologists in this field (for a selection of representative examples, see Bertelli and Giannini 2005, Pérez-Emán 2005, García-Moreno et al 2006, Ribas et al 2007, Chaves et al 2009, Parra et al 2009, Tello et al 2009). These outstanding publications are the result not only of improved academic training, but also of increased opportunities for international and inter-institutional collaboration (Naranjo 2008).…”