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Family Names

           There are two, three, four, or five families in the order Falconiformes, or diurnal raptors, depending on one’s taxonomic views, but there is a growing consensus that the group is not monophyletic. Indeed, Remsen et al. (2008) commented that they were unaware of any genetic data supporting the monophyly of the order except for those of Sibley and Ahlquist (1990). The morphological analyses of Livezey and Zusi (2007) and Mayr and Clarke (2003) appear to provide the primary evidence for monophyly, and these similarities may be the result of convergence (Remsen et al. op cit. 2008).

          There is general agreement as to the inclusion of the families Accipitridae (263 species of hawks, eagles, harriers, and kites) and Sagittariidae (one species, the Secretarybird), and the Osprey (Pandion haliaetus) has usually been placed in its own family, Pandionidae (Brown and Amadon 1968, American Ornithologists’ Union 1983, del Hoyo et al. 1994, Ferguson-Lees and Christie 2001). However, some important recent references have merged the Osprey as a subfamily within the Accipitridae (Stresemann and Amadon 1979, Sibley and Monroe 1990, Dickinson 2003, American Ornithologists’ Union 2008). Based on the nucleotide sequences of the mitochondrial cytochrome-b gene, Wink and Sauer-Gürth (2004) concluded that Pandion is sufficiently distinct to be maintained in its own family, and this was also confirmed by Lerner and Mindell (2005), based on a study using both mitochondrial and nuclear DNA sequences. Their recommendation is followed here.

          There has been perennial lack of consensus among taxonomists over the evolutionary relationships of the family Cathartidae, which includes seven species of New World vultures and condors. During the 1990s, most authorities placed this enigmatic, but clearly monophyletic, group near the storks (Ciconiiformes), based on a variety of molecular, behavioral, and morphological characters (Ligon 1967, König 1982, Rea 1983, Jacob 1983, Sibley and Ahlquist 1990, Avise et al. 1994, Seibold and Helbig 1995, Wink et al. 1998). However, this treatment has been strongly disputed by more recent studies (e.g., Griffiths 1994, Cracraft et al. 2004, Ericson et al. 2006, Gibb et al. 2007, Slack et al. 2007, Livezey and Zusi 2007, Hackett et al. 2008), drawing from both molecular and morphological evidence, and the American Ornithologists’ Union Committee on Classification and Nomenclature recently recommended that the Cathartidae again be included in the Falconiformes (Banks et al. 2007). The AOU’s South American Classification Committee regards the placement of the cathartids as still uncertain and is considering a proposal to place them in a separate order, Cathartiformes (Remsen et al. 2008).

          Judging from a preponderance of genetic studies (van Tuinen et al. 2000, Fain and House 2004, Ericson et al. 2006, Mayr and Clarke 2003, Gibb et al. 2007), the family Falconidae (59 species of true falcons, forest falcons, falconets, caracaras, and several related forms) does not appear to be closely related to the other accipitriform families. In fact, the recent comprehensive analysis of ~32 kilobases of nuclear DNA gene sequences of all major extant avian groups by Hackett et al. (2008) indicated that falcons forms a sister clade to songbirds (Passeriformes) and parrots (Psittaciformes), quite distant from the Accipitridae.

          Despite their apparent polyphyletic origins, the diurnal birds of prey share many behavioral and life history traits, and the main conservation issues are similar for these families, justifying their inclusion in the GRIN project.

          The genera/number of species included in GRIN are as follows:

          Family Cathartidae (5/7):
               Cathartes: 3
               Coragyps: 1
               Gymnogyps: 1
               Sarcoramphus: 1
               Vultur: 1

          Family Pandionidae (1/1):
               Pandion: 1

          Family Accipitridae (65/262):
               Accipiter: 52
               Aegypius: 1
               Aquila: 11
               Aviceda: 5
               Busarellus: 1
               Butastur: 4
               Buteo: 26
               Buteogallus: 5
               Chelictinia: 1
               Chondrohierax: 2
               Circaetus: 6
               Circus: 16
               Dryotriorchis: 1
               Elanoides: 1
               Elanus: 4
               Erythrotriorchis: 2
               Eutriorchis: 1
               Gampsonyx: 1
               Geranospiza: 1
               Gypaetus: 1
               Gypohierax: 1
               Gyps: 8
               Haliaeetus: 8
               Haliastur: 2
               Hamirostra: 1
               Harpagus: 2
               Harpia: 1
               Harpyhaliaetus: 2
               Harpyopsis: 1
               Helicolestes: 1
               Henicoperrnis: 1
               Hieraaetus: 5
               Ichthyophaga: 2
               Ictinaetus: 1
               Ictinia: 2
               Kaupifalco: 1
               Leptodon: 2
               Leucopternis: 10
               Lophoaetus: 4
               Lophoictinia: 1
               Lophotriorchis 1
               Machaeramphus: 1
               Megatriorchis: 1
               Melierax: 3
               Micronisus: 1
               Milvus: 2
               Morphnus: 1
               Necrosyrtes: 1
               Neophron: 1
               Nisaetus: 9
               Parabuteo: 1
               Percnohierax: 1
               Pernis: 4
               Pithecophaga: 1
               Polemaetus: 1
               Polyboroides: 2
               Rostrhamus: 1
               Rupornis: 1
               Sarcoramphus: 1
               Spilornis: 13
               Spizaetus: 4
               Stephanoaetus: 1
               Terathopius: 1
               Torgos: 1
               Trigonoceps: 1
               Urotriorchis: 1

          Family Pandionidae (1/1):
               Pandion: 1

          Family Sagittariidae (1/1):
               Sagittarius: 1

          Family Falconidae (11/59):
               Caracara: 3
               Daptrius 1
               Falco: 39
               Herpetotheres: 1
               Ibycter: 1
               Micrastur: 7
               Microhierax: 5
               Milvago: 2
               Phalcoboenus: 4
               Polihierax: 2
               Spiziapteryx: 1
Literature Cited:

American Ornithologists' Union. 1983. Check-list of North American birds, 6th edition. American Ornithologists' Union, Washington, D.C.

American Ornithologists' Union. 1998. Check-list of North American birds. 7th edition. American Ornithologists' Union, Washington, D.C.

Avise, J. C., W. S. Nelson, and C. G. Sibley. 1994. DNA sequence support for a close phylogenetic relationship between some storks and New World vultures. Proceedings of the National Academy of Sciences 91:5173-5177.

Banks, R.C., R.T. Chesser, C. Cicero, J.L. Dunn, A.W. Kratter, I.J. Lovette, P.C. Rasmussen, J.V. Remsen, Jr., J.D. Rising, and D.F. Stotz. 2007. Forty-eighth supplement to the American Ornithologists’ Union Check-list of North American Birds. Auk 124:1109-1115.

Brown, L. H., and D. Amadon. 1968. Eagles, hawks, and falcons of the world. Country Life Books, London.

Cracraft, J., F.K. Barker, M. Braun, J. Harshman, G.J. Dyke, J. Feinstein, S. Stanley, A. Cibois, P. Schikler, P. Beresford, et al. 2004. Phylogenetic relationships among modern birds (Neornithes): toward an avian tree of life. Pp. 468-489 in J. Cracraft and M.J. Donoghue (eds.), Assembling the tree of life. Oxford University Press, Oxford, UK.

del Hoyo, J., A. Elliott, and J. Sargatal (eds.). 1994. Handbook of birds of the world. Vol 2. New World vultures to guineafowl. Lynx Edicions, Barcelona, Spain.

Dickinson, E.C. (ed.). 2003. The Howard and Moore complete checklist of the birds of the world. 3rd edition. Princeton University Press, Princeton, NJ.

Ericson, P.G.P., C.L. Anderson, T. Britton, A. Elzanowski, U.S. Johansson, M. Kallerrsjo, J.I. Ohlson, T.J. Parsons, D. Zuccon, and G. Mayr. 2006. Diversification of Neoaves: integration of molecular sequence data and fossils. Biology Letters 2:543-547.

Fain, M.G., and P. Houde. 2004. Parallel radiations in the primary clades of birds. Evolution 58:2558-2573.

Ferguson-Lees, J., and D. A. Christie. 2001. Raptors of the world. Houghton Mifflin, Boston.

Gibb, G.C., R.T. Kardailsky, R.T. Kimball, E.L. Braun, and D. Penny. 2007. Mitochondrial genomes and avian phylogeny: complex characters and resolvability without explosive radiations. Molecular Biology Evolution 24:269-280.

Griffiths, C.S. 1994. Monophyly of the Falconiformes based on syringeal morphology. Auk 111:787-805.

Hackett, S.J., R.T. Kimball, S. Reddy, R.C.K. Bowie, E.L. Braun, M.J. Braun, J.L. Chojnowski, W.A. Cox, K.-L. Han, J. Harshman, C.J. Huddleston, B.D. Marks, KJ.J. Miglia, W.S. Moore, F.H. Shelcon, D.W. Steadman, C.C. Witt, and T. Yuri. 2008. A phylogenetic study of birds reveals their evolutionary history. Science 320:1763-1768.

Jacob, J. 1983. Zur systematischen Stellung von Vultur gryphus (Cathartiformes). Journal für Ornithologie 124:83-86.

König, C. 1982. Zur systematischen Stellung der der Neuweltgeier (Cathartidae). Journal für Ornithologie 123:259-267.

Lerner, H.R., and D.P. Mindell. 2005. Phylogeny of eagles, Old World vultures, and other Accipitridae based on nuclear and mitochondrial DNA. Molecular Phylogenetics and Evolution 37:327-346.

Ligon, J.D. 1967. Relationships of the cathartid vultures. University of Michigan, Museum of Zoology, Occasional Papers no. 651.

Livezey, B.C., and R.L. Zusi. 2007. Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion. Zoological Journal of the Linneaean Society 149:1-95.

Mayr, G., and J. Clarke. 2003. The deep divergences of neornithine birds: a phylogenetic analysis of morphological characters. Cladistics 19:527-553.

Poole, A.F. 1994. Family Pandionidae (Osprey). Pp. 42-51 in J. del Hoyo, A. Elliott, and J. Sargatal (eds.). Handbook of birds of the world. Vol 2. New World vultures to guineafowl. Lynx Edicions, Barcelona, Spain.

Rea, A.M. 1983. Cathartid affinities: a brief overview. Pp. 26-54 in S.R. Wilbur and J.A. Jackson (eds.), Vulture biology and management. University of California Press, Berkeley, CA.

Remsen, J.V., Jr. (Chair). 2008. A classification of the bird species of South America. (Part 2). South American Classification Committee, American Ornithologists’ Union, Baton Rouge, LA.

Seibold, I., and A.J. Helbig. 1995. Zur systematischen Stellung des Fischadlers Pandion haliaetus nach mitochondriellen DNA-Sequenzen. Vogelwelt 116:209-217.

Sibley, C.G., and J.E. Ahlquist. 1990. Phylogeny and classification of birds. Yale University Press, New Haven, CT.

Sibley, C.G., and B.L. Monroe, Jr. 1990. Distribution and taxonomy of birds of the world. Yale University Press, New Haven, CT.

Slack, K.E., F. Delsuc, P.A. McClenachan, U. Arnason, and D. Penny. 2007. Resolving the root of the avian mitogenic tree by breaking up long branches. Molecular Phylogenetics and Evolution 42:1-13.

Stresemann, E., and D. Amadon. 1979. Falconiformes. Pp. 271-425 in E. Mayr and G.W. Cottrell (eds.), Check-list of birds of the world. Vol. 1. 2nd edition. Museum of Comparative Zoology, Cambridge, MA.

Van Tuinen, M., C.G. Sibley, and S.B. Hedges. 2000. The early history of modern birds inferred from DNA sequences of nuclear and mitochondrial ribosomal genes. Molecular Biology & Evolution 17:451-457.

Wink, M. 1995. Phylogeny of Old and New World vultures (Aves: Accipitridae and Cathartidae) inferred from nucleotide sequences of the mitochondrial cytochrome b gene. Verlag der Zeitschrift für Naturforschung 50c:868-882.

Wink, M., and H. Sauer-Gürth. 2004. Phylogenetic relationships in diurnal raptors based on nucleotide sequences of mitochondrial and nuclear marker genes. Pp. 483-498 in R.D. Chancellor and B.-U. Meyburg (eds.), Raptors worldwide. World Working Group on Birds of Prey and Owls, Berlin, and MME/BirdLife Hungary, Budapest.

Wink, M., L. Siebold, F. Lotfikham, and W. Bednarek. 1998. Molecular systematics of Holarctic raptors (Order Falconiformes). Pp. 29-48 in R.D. Chancellor, B.-U. Meyburg, and J.J. Ferrero (eds.), Holarctic birds of prey. ADENEX and World Working Group on Birds of Prey, Calamonte, Spain.

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