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Combined Phylogenetic Analysis of a New North American Fossil Species Confirms Widespread Eocene

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Combined Phylogenetic Analysis of a New North American Fossil Species Confirms Widespread Eocene Zoological Journal of the Linnean Society, 2009, 157, 586–611. With 9 figures Combined phylogenetic analysis of a new North American fossil species confirms widespread Eocene distribution for stem rollers (Aves, Coracii)zoj_550 586..611 JULIA A. CLARKE1,2,3,4*, DANIEL T. KSEPKA1,2,3, N. ADAM SMITH1,4 and MARK A. NORELL3 1Department of Marine, Earth and Atmospheric Sciences, North Carolina State University, Campus Box 8208, Raleigh, NC 27695-8208, USA 2Department of Paleontology, North Carolina Museum of Natural Sciences, 11 West Jones St, Raleigh, NC 27601-1029, USA 3Division of Paleontology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA 4New Address: Department of Geological Sciences, Jackson School of Geosciences, The University of Texas, PO Box B University Station, Austin, TX 78713-8902, USA Received 30 August 2008; accepted for publication 17 November 2008 We report a nearly complete skeleton of a new species of stem roller (Aves, Coracii) from the early Eocene Green River Formation of North America. The new species is most closely related to two species-depauperate lineages, Coraciidae (rollers) and Brachypteraciidae (ground rollers), that form a monophyletic crown clade (Coracioidea) with an exclusively Old World extant distribution. Phylogenetic analysis utilizing a matrix of 133 morphological characters and sequence data from three genes (RAG-1, c-myc, and ND2) identifies the new species as a stem member of the Coracii more closely related to the crown clade than the only previously identified New World taxon, Primobucco mcgrewi. The phylogenetic placement of the new species and Primobucco mcgrewi indicates a widespread northern hemisphere distribution in the Eocene with subsequent restriction to Africa, Madagascar, Australia, and temperate to tropical parts of Europe and Asia. It provides evidence of further ecological diversity in early stem Coracii and convergence on crown morphologies. The new species contributes to mounting evidence that extant distributions for major avian subclades may be of comparatively recent origin. Further late Palaeogene sampling is needed to elucidate potential drivers for shifting avian distributions and disappearance of Coracii from North America. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157, 586–611. doi: 10.1111/j.1096-3642.2009.00550.x ADDITIONAL KEYWORDS: avian evolution – biogeography – Cenozoic – combined analysis – Green River Formation – phylogeny. INTRODUCTION nantly terrestrial Brachypteraciidae are endemic to Madagascar, whereas the more arboreal ‘true’ rollers Rollers (Coraciidae) and ground rollers (Brac- have an extant distribution that spans Europe, Africa, hypteraciidae) are medium-sized birds with colourful south Asia, and Australia (Fry, Fry & Harris, 1992; plumage, large heads, and recurved beaks. The Langrand, 2001; Fig. 1). These two taxa formed the common name roller is derived from the characteristic clade Coracioidea of Cracraft (1971, 1981). The Lep- acrobatic display flights of ‘true’ rollers (Coraciidae), tosomidae (cuckoo-rollers) are represented by a single although the ground rollers do not exhibit this behav- predominately arboreal species from Madagascar and iour (Whitfield, 1988). The five species of predomi- the Comoro Islands (Langrand, 2001) and have at times been considered closely related to the Coracio- *Corresponding author. E-mail: [email protected] idea, largely following the early work of Sclater 586 © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157, 586–611 NEW SPECIES FROM THE EOCENE GREEN RIVER FORMATION 587 Figure 1. Map showing the distribution of extant and extinct Coracii. (1865). However, this relationship has been debated 2006; Livezey & Zusi, 2007; Hackett et al., 2008). (e.g. Mayr & Amadon, 1951; Wetmore, 1960; Maurer More recently, the clade including only extinct stem & Raikow, 1981; Mayr, 1998; Cracraft et al., 2004) taxa more closely related to Coracioidea than to any and is unsupported in most recent analyses other extant lineage was referred to as ‘Coracii- (e.g. Sibley & Ahlquist, 1990; Mayr & Clarke, 2003; formes sensu stricto’ by Mayr (1998). Given that the Mayr, Manegold & Johansson, 2003; Mayr, Mourer- taxon Coraciiformes has consistently included an Chauviré & Weidig, 2004; Ericson et al., 2006; array of other ‘higher land birds’ as well as rollers Hackett et al., 2008; although see Livezey & Zusi, over the past 100 years (e.g. Forbes, 1884; Für- 2006, 2007 and response by Mayr, 2008). bringer, 1888; Stresemann, 1959; Sibley, Ahlquist & The taxon name ‘Coraciiformes’ was used by Monroe, 1988; Sibley & Ahlquist, 1990) and is cur- Fürbringer (1888) for Coraciidae, Leptosomidae, rently so used by most contemporary references (e.g. Caprimulgiformes, and Strigiformes, and in subse- Höfling & Alvarenga, 2001; Johansson & Ericson, quent classifications, it has been applied to distinct 2003; Cracraft et al., 2004; Livezey & Zusi, 2007; sets of ‘higher landbird’ taxa. The proposed compo- Hackett et al., 2008), it seems best to reserve this nent taxa have varied significantly, and which taxa taxon name for this more inclusive clade in keeping might comprise a monophyletic core with the Coraci- with its prior use. We instead recommend the avail- idae and Brachypteraciidae remains debated (Mayr able taxon name ‘Coracii’, which has been consis- & Clarke, 2003; Mayr et al., 2003; Ericson et al., tently used at the ‘subordinal’ level (e.g. Wetmore & 2006; Livezey & Zusi, 2007; Hackett et al., 2008). Miller, 1926; Stresemann, 1959; Wetmore, 1960; Proposed parts of this clade have included hornbills Cracraft, 1971; Maurer & Raikow, 1981; Burton, (Bucerotiformes), hoopoes (Upupiformes), trogons 1984; Sibley et al., 1988; Sibley & Ahlquist, 1990; (Trogonidae), motmots (Momotidae), todies (Todidae), Johansson & Ericson, 2003), to be phylogenetically bee-eaters (Meropidae), and kingfishers (Alcedinidae) defined for the stem clade including all taxa more (e.g. Gadow, 1892; Stresemann, 1959; Wetmore, closely related to Coracioidea than to its nearest 1960; Cracraft, 1981; Maurer & Raikow, 1981; Sibley outgroup [equivalent contents to ‘Coraciiformes & Ahlquist, 1990; Mayr et al., 2004; Ericson et al., sensu stricto’ of Mayr (1998); with external specifiers © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157, 586–611 588 J. A. CLARKE ET AL. amongst the other major lineages of traditional of Quercy, France (Feduccia, 1999), but this place- coraciiform and piciform birds]. ment was recently found to be unsupported (Mayr & Wetmore & Miller (1926) were apparently the first Mourer-Chauviré, 2000). Kirchman et al. (2001) cited to use ‘Coracii’ instead of ‘Coraciae’ while standardiz- undescribed fossils attributed to the Brachypteraci- ing subordinal endings across Aves and recognizing idae from the middle Eocene of Europe as a calibra- restricted contents relative to Fürbringer (1888) and tion point for the Coraciidae/Brachypteraciidae split other prior authors. ‘Coracii’ of Wetmore & Miller in divergence estimates for the timing of ground roller (1926), unlike prior usages of the taxon name ‘Cora- diversification. These authors concluded that, given ciae’, included rollers but no other ‘higher landbird’ the antiquity of the lineage inferred from these fossil taxa. The name ‘Coraciae’ was later resurrected for a calibrations, there was a notable lack of sequence ‘parvclass’ including Coraciiformes (e.g. Sibley et al., divergence within Brachypteraciidae relative to other 1988; Sibley & Ahlquist, 1990), although this taxon groups of birds. However, they did note that this name does not appear to be in broad use. referral could be questionable and proposed that the We use ‘Coracioidea’ consistent with its originally fossils could be found to be outside the roller crown recognized contents, including Coraciidae and Brac- clade. Such a placement was indeed confirmed for the hypteraciidae and excluding Leptosomidae (Cracraft, middle Eocene Messel roller referenced by Kirchman 1971), but recommend redefining the term phyloge- et al. (2001) now named Eocoracias brachyptera netically as the most recent common ancestor of (Mayr & Mourer-Chauviré, 2000, 2003). Coraciidae and Brachypteraciidae and all of its Mayr & Mourer-Chauviré (2000) described new descendents. The monophyly of this crown clade has material of Geranopterus alatus Milne-Edwards been supported by morphological (Mayr et al., 2003, (1892) and a new species (Geranopterus milneed- 2004) and molecular sequence data (e.g. Kirchman wardsi) from deposits within the Quercy fissure fills et al., 2001; Cracraft et al., 2004; Ericson et al., 2006; (dated to approximately 35 Myr; Legendre & Lévêque, Hackett et al., 2008). Following the usage recom- 1997), identifying these as part of the coraciiform stem mended here, whether Coracioidea includes the fossil lineage. More recently, further isolated material has taxon Geranopteridae hinges on whether it is recov- been referred to Geranopteridae from these late ered to be part of the crown clade (see Discussion). Eocene deposits (Mourer-Chauviré & Sigé, 2006). This usage is distinct from that of Mayr & Mourer- Intriguingly, Mourer-Chauviré (1999) reassigned a Chauviré (2000) who applied this taxon name to a tarsometatarsus described as a fossil jacana by Mlík- more inclusive clade (crown clade + Geranopteridae). ovsky (Nupharanassa bohemica Mlíkovsky, 1999) to An array of fossil taxa from Europe
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