Aves, Coraciiformes SENSU STRICTO), with first Records from Europe

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Aves, Coraciiformes SENSU STRICTO), with first Records from Europe Journal of Systematic Palaentology 2 (1): 1–12 Issued 11 March 2004 DOI: 10.1017/S1477201903001093 Printed in the United Kingdom C The Natural History Museum Osteology and systematic position of the Eocene Primobucconidae (Aves, Coraciiformes SENSU STRICTO), with first records from Europe Gerald Mayr Forschungsinstitut Senckenberg, Division of Ornithology, Senckenberganlage 25, 60325 Frankfurt am Main, Germany Cecile´ Mourer-Chauvire´ UMR 5125 Pal´eoenvironnements et Pal´eobiosph`ere, Centre des Sciences de la Terre, Universit´eClaude Bernard-Lyon 1, 27–43 Boulevard du 11 Novembre, 69622 Villeurbanne Cedex, France Ilka Weidig Forschungsinstitut Senckenberg, Division of Ornithology, Senckenberganlage 25, 60325 Frankfurt am Main, Germany SYNOPSIS The first complete skeletons of the early Eocene avian taxon Primobucconidae Brodkorb, 1970 are described and the osteology and systematic position of these poorly known birds are revised. New specimens from Germany (Messel) and France (Cond´e-en-Brie) are the first Old World records of this taxon. From Messel, two new species, Primobucco perneri sp. nov. and P. frugilegus sp. nov., are described. Cladistic analysis of 36 morphological characters from 14 ingroup taxa placed the Primobucconidae in an unresolved polytomy including fossil and extant rollers (Coraciiformes sensu stricto). Derived characters are presented that support this classification. The Primobucconidae is yet another taxon that exemplifies the great similarity between the early Eocene avifaunas of North America and Europe. It further constitutes the first record of stem group rollers in the New World. Both known specimens of P. frugilegus are preserved with seeds in the area of the stomach, which may indicate that stem group rollers had a more generalised diet than their extant relatives. KEY WORDS Fossil birds, Rollers, Phylogeny, Messel, Green River Formation, Paris Basin Contents Introduction 2 Material and methods 2 Terminology 2 Phylogenetic analysis 2 Institutional abbreviations 3 Systematic descriptions 3 CORACIIFORMES sensu stricto (see Mayr, 1998b)3 PRIMOBUCCONIDAE Feduccia & Martin, 1976 3 Genus PRIMOBUCCO Brodkorb, 1970 4 Primobucco mcgrewi Brodkorb, 1970 4 Primobucco perneri sp. nov. 5 Primobucco frugilegus sp. nov. 7 Primobucconidae gen. et sp. indet. 7 Results of phylogenetic analysis 8 Discussion 8 Acknowledgements 9 References 9 Appendix 1 11 Appendix 2 12 2 G. Mayr ET AL. Introduction conidae from Messel and an isolated tarsometatarsus from the Eocene of France. For the first time complete skeletons The avian species Primobucco mcgrewi was described by of the Primobucconidae are described and the phylogenetic Brodkorb (1970) on the basis of an isolated wing from the relationships of this taxon are evaluated in a cladistic ana- Lower Eocene Green River Formation (Wyoming, USA). It lysis. was considered to be the earliest representative of the Neo- tropical puffbirds (Bucconidae, Galbulae), which constitute part of the traditional (e.g. Wetmore 1960) Piciformes (wood- Material and methods peckers and allies: (Simpson & Cracraft 1981; Swierczewski & Raikow 1981; Johansson & Ericson 2003; Mayr et al. Terminology 2003). Feduccia & Martin (1976) later erected the taxon Primobucconidae for P. mcgrewi and four other early Eo- The traditional Coraciiformes sensu Wetmore (1960) are cene genera that they assumed to be the ‘dominant small probably a polyphyletic taxon, since the Upupiformes and perching birds of the Eocene of North America’ (Feduccia & Bucerotiformes may be more closely related to piciform Martin 1976: 101). Olson & Feduccia (1979) further repor- birds, while cuckoo-rollers (Leptosomidae) are probably ted a record of the Primobucconidae from the Lower Eocene more closely related to ‘non-coraciiform’ birds (e.g. Mayr London Clay of England, but this identification was disputed et al. 2003). Thus, following Mayr (1998b), the term Coracii- by Harrison (1982) who instead assigned another specimen formes sensu stricto (Coraciiformes s.s.) is used here for the from the same deposits to the Primobucconidae. clade that includes the Eocoraciidae, Geranopteridae, Cora- More recently, however, it has been shown that the ciidae and Brachypteraciidae, the term Alcediniformes is Primobucconidae, as established by Feduccia & Martin used for the clade that includes the Meropidae, Alcedinidae, (1976), are a polyphyletic assemblage of birds which be- Momotidae and Todidae, while the term Upupiformes is used long to different higher taxa, such as stem group mousebirds for the clade that includes the Upupidae and Phoeniculidae. (Coliiformes: see Houde & Olson 1992; Mayr & Peters 1998) Osteological terminology follows Baumel & Witmer (1993). and stem group parrots (Psittaciformes: see Mayr 1998a, 2002). The only taxon currently recognised in the Primobuc- conidae is Primobucco mcgrewi (Houde & Olson 1989) and Phylogenetic analysis the holotypic wing is still the only specimen of this species The Primobucconidae have been considered, so far, to be that has been described. However, Houde & Olson (1989: closely related to either the Galbulae (Brodkorb 1970; 2032) noticed that the holotype of P. mcgrewi ‘compares Feduccia & Martin 1976) or to rollers (Houde & Olson 1989). best with one of the newly obtained skeletons from the Green Our analysis covers these taxa and, based on the existing River Formation that represents an undescribed, facultatively phylogenies of the ‘higher land bird assemblage’ (e.g. Sibley zygodactyl (i.e. the fourth toe can be turned backwards), & Ahlquist 1990; Johansson et al. 2001; Mayr et al. 2003), roller-like bird’. These authors further indicated (p. 2030) taxa that are presumed to be closely related forms. Outgroup that P. mcgrewi ‘may belong with roller-like birds similar to comparisons were made with the Musophagidae (turacos) the Madagascan Atelornithidae [= Brachypteraciidae]’. and the Caprimulgidae (nighthawks). Rollers (Coraciiformes sensu stricto), including the ex- For the phylogenetic analysis, skeletons of the ex- tant true rollers (Coraciidae) and the ground rollers (Bra- tant taxa listed below were examined in the collections of chypteraciidae) have a poor early Tertiary fossil record. the Forschungsinstitut Senckenberg, the Museum´ National Early Tertiary stem group representatives of rollers were only d’Histoire Naturelle, Paris and the Museum fur¨ Naturkunde, known, so far, from the Middle Eocene of Messel, Germany Berlin. (Eocoraciidae) and the Upper Eocene of the Quercy fissure fillings in France (Geranopteridae: Mayr & Mourer-Chauvire´ Musophagidae: Corythaixoides, Crinifer, Musophaga, 2000). Tauraco. Molecular studies suggest that rollers are the sister taxon Caprimulgidae: Caprimulgus, Chordeiles. of the alcediniform birds, which include bee-eaters (Merop- idae), kingfishers (Alcedinidae), motmots (Momotidae) and Galbulidae: Galbula. todies (Todidae: Sibley & Ahlquist 1990, Espinosa de los Bucconidae: Chelidoptera, Monasa, Notharchus. Monteros 2000, Johansson et al. 2001, Mayr et al. 2003). Ramphastidae: Megalaima, Psilopogon, Pteroglossus, The earliest alcediniform bird described so far was also Ramphastos, Selenidera, Trachyphonus. from Messel (Mayr 1998b); other early Tertiary Alcedini- Indicatoridae: Indicator. formes are known from the Upper Eocene of France and the Picidae: Colaptes, Dendrocopos, Dendropicos, Jynx, Picus. Oligocene of Switzerland and North America (Olson 1976; Coraciidae: Coracias, Eurystomus. Mourer-Chauvire´ 1985). An as yet undescribed ‘kingfisher- Brachypteraciidae: Atelornis, Geobiastes, Uratelornis. like bird’ from the Green River Formation figured in Grande Meropidae: Merops. (1980: fig. III.19) and Feduccia (1999: 335) belongs to the Alcedinidae: Alcedo, Ceryle, Dacelo, Halcyon, Ispidina. Upupiformes (hoopoes, wood-hoopoes and allies), as indic- Momotidae: Momotus. ated by, e.g. the very long hallux and the derived morphology of the carpometacarpus and distal tarsometatarsus. Todidae: Todus. A revision of the avifauna of the Green River Formation Leptosomidae: Leptosomus. by one of us (I.W.) led to the identification of new specimens Upupidae: Upupa. of P. mcgrewi from the type locality, which are presented in Phoeniculidae: Phoeniculus. this study. We further describe two new taxa of the Primobuc- Bucerotidae: Bucorvus, Bycanistes, Tockus, Penelopides. Osteology and systematics of Eocene Primobucconidae (Aves) 3 A total of 36 characters from 16 terminal taxa (see teridae, which touches the jugal bars. The ‘head’ (Cracraft character matrix in Appendix 2) were coded for a parsimony 1968: fig. 2) of the os lacrimale (SMF-ME 3507a, SMF-ME analysis using PAUP 3.1 (Swofford 1993). All characters 3794) is large, as in extant rollers, Alcedinidae and Galbulae. were coded as unordered. The shortest tree was found using the branch-and-bound search option. The consistency index Vertebrae. The number of praesacral vertebrae appears to (CI), retention index (RI) and rescaled consistency index have been about 19, as in most ‘higher land birds’ (pers. obs.). (RC) were calculated. The robustness of the tree was tested The incisura fossae of the atlas is dorsally open (visible in with a bootstrap analysis of 1000 replicates using the same Primobucco mcgrewi, USNM 336284). There are seven free settings as in the primary analysis. tail vertebrae (SMNK.PAL.404a+b). The pygostyle bears a cranio-caudally wide processus dorsalis, which is also found in extant rollers and alcediniform birds. Institutional Abbreviations Coracoid. The extremitas omalis of the coracoid (SMF-
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