Project 510: B. C. LIVEZEY, 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 Linnean Society. 149 (1):1-95.
Specimen: Rhea (unvouchered)

Abstract

This work serves as a compendium of anatomical resources upon which a companion phylogenetic analysis of Aves and related Theropoda (Avialae) was based (Livezey and Zusi 2006). Following a brief historical overview of avian anatomy and avian systematics, the rich published literature pertinent to these topics is classified chronologically and geographically. The former also was organized with respect to eras of predominant methodologies in avian systematics, with an emphasis on recent paleontological finds bearing importantly on the origins of modern birds (Neornithes). This was followed by an exposition on the theoretical and abstract underpinnings of morphological characterization for purposes of phylogenetic reconstructions (to be published separately), with aspects of analysis for phylogenetic inference (e.g., tactics for employment of currently available software, ordering, criteria for optimization of trees) considered elsewhere.

The principal contribution of this exposition is a listing of characters and states manifesting what was inferred to hold promise with respect to phylogenetic or historical signal. In total, 2,954 anatomical characters were defined—2,451 osteological, 256 myological, and 247 miscellaneous—of which 981 (approximately one-third) were multiple-state (i.e., comprised three or more states), the latter including 537 characters treated as ordered. Bibliographic provenance for characters was provided, where possible, but exact equivalence of characters and states among workers was seldom feasible. In many cases, previously published characters were listed on the grounds that these pertained most closely to the structure or complex at hand in the present study, and that such listings provided at least a historical grasp of the magnitude of prior usages of a given character. We also summarized, to the extent feasible, previously published characters for which inclusion in the present work was judged to be unreliable or lacking sufficient clarity.

In addition to the descriptions of characters, a limited series of figures are provided, in no small part to ameliorate the challenges posed by new terms and formal anatomical nomenclature. We are adherents to nomenclatural formalism (Livezey and Zusi 2001) in anatomical contexts sensu the ICAAN (International Committee on Avian Anatomical Nomenclature; Baumel 1993). I.e., we consider that characters and states—implicitly proposals of homology—warrant clarity with respect to surrounding text in the same sense that binomial taxa—i.e., as hypotheses of historical lineages—are subject to formal conventions.

Finally, literature cited herein is listed, as a work of this kind is impossible without access to the wealth of information and insight provided by such a resource. As this literature is integrated by citation with the descriptions of characters, it is hoped that the bibliography will lessen the challenges posed by a deep, multilingual, and variably technical literature for systematists using these descriptions. The dimension of the character matrix also led us to enclose a CD of the data set in the present work to assist those seeking to improve, append, or refine our efforts.

A phylogenetic (cladistic) analysis of these data will appear separately (Livezey and Zusi 2006). Soon thereafter a collaboration with an unparalleled compilation of molecular data (i.e., DNA sequences) and confirmatory paleontological data is planned to conclude with the publication of a total-evidence analysis of avian phylogeny under the auspices of the NSF “Tree of Life” program, one to encompass extant birds (Neornithes), avian relatives from the Mesozoic (outgroups), and nonavialian fossil taxa from the Mesozoic (“deep” outgroups).



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Article DOI: 10.1111/j.1096-3642.2006.00293.x

Project DOI: 10.7934/P510, http://dx.doi.org/10.7934/P510
This project containsMatrices
  • 130 Media
  • 1 Matrix
  • 5 Documents
  • 188 Taxa
  • 128 Specimens
  • 2954 Characters
Total size of project's media files: 46.16M

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Total scored cells: 433314
Total media associated with cells: 0
Total labels associated with cell media: 0
Characters
Total characters: 2954
Total characters with associated media: 0
Total characters with media with labels: 0
Total character states: 8358
Total character states with associated media: 0
Total character states with media with labels:0
Total unordered/ordered characters:2954/0
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    Taxonomic Overview for Matrix 'M1228' (188 Taxa)

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    [1] Non-Archosauromorpha
    Taxon name last Modified on 12/05/13
    284912610500004
    [2] Crocodylia
    Taxon name last Modified on 03/16/17
    280814814600004
    [3] Ornithischia
    Taxon name last Modified on 03/16/17
    267529327900004
    [4] Sauropodomorpha
    Taxon name last Modified on 03/16/17
    264531930900004
    [5] Herrerasaurus
    Taxon name last Modified on 03/16/17
    237358158100004
    [6] Dilophosaurus
    Taxon name last Modified on 12/05/13
    247947647500004
    [7] Abelisauridae
    Taxon name last Modified on 03/16/17
    270924624500004
    [8] Coelophysis
    Taxon name last Modified on 12/05/13
    240555954900004
    [9] Ceratosaurusia
    Taxon name last Modified on 12/05/13
    249146846300004
    [10] Allosauroidea
    Taxon name last Modified on 12/05/13
    216383479100004
    [11] Tyrannosauridae
    Taxon name last Modified on 03/16/17
    222274873200004
    [12] Compsognathus
    Taxon name last Modified on 12/05/13
    248347347100004
    [13] Sinosauropteryx
    Taxon name last Modified on 12/05/13
    274920620500004
    [14] Caudipteryx
    Taxon name last Modified on 12/05/13
    255240740200004
    [15] Protoarchaeopteryx
    Taxon name last Modified on 12/05/13
    277517917900004
    [16] Oviraptorosauria
    Taxon name last Modified on 03/16/17
    227769267700004
    [17] Ornithomimosaurus
    Taxon name last Modified on 12/05/13
    227270368200004
    [18] Struthiomimus
    Taxon name last Modified on 03/16/17
    274021621400004
    [19] Gallimimus
    Taxon name last Modified on 12/05/13
    273322322100004
    [20] Pelecanimimus
    Taxon name last Modified on 12/05/13
    276718718700004
    [21] Therizinosauroidea
    Taxon name last Modified on 03/16/17
    243852651600004
    [22] Troodon
    Taxon name last Modified on 12/05/13
    226469469000004
    [23] Saurornithoides
    Taxon name last Modified on 12/05/13
    279915515500004
    [24] Dromaeosaurus
    Taxon name last Modified on 03/16/17
    213484482000004
    [25] Velociraptor
    Taxon name last Modified on 12/05/13
    253641841800004
    [26] Deinonychus
    Taxon name last Modified on 12/05/13
    263831631600004
    [27] Alvarezsauroidea
    Taxon name last Modified on 12/05/13
    232467763000004
    [28] Enantiornithes
    Taxon name last Modified on 03/16/17
    246851748600004
    [29] Archaeopteryx
    Taxon name last Modified on 12/05/13
    16601297129400004
    [30] Confuciusornis
    Taxon name last Modified on 12/05/13
    17831175117100004
    [31] Hesperornis
    Taxon name last Modified on 12/05/13
    243452452000004
    [32] Ichthyornis
    Taxon name last Modified on 12/05/13
    17751184117900004
    [33] Rahonavis
    Taxon name last Modified on 12/05/13
    284610810800004
    [34] Apsaravis
    Taxon name last Modified on 12/05/13
    253342342100004
    [35] Lithornis
    Taxon name last Modified on 12/05/13
    15771384137700004
    [36] Aepyornithidae
    Taxon name last Modified on 03/16/17
    11921765176200004
    [37] Dinornithiformes
    Taxon name last Modified on 03/16/17
    11911795176300004
    [38] Struthio
    Taxon name last Modified on 03/16/17
    4072550254700004
    [39] Rhea
    Taxon name last Modified on 03/16/17
    3892567256500004
    [40] Casuarius
    Taxon name last Modified on 03/16/17
    3842571257000004
    [41] Dromaius
    Taxon name last Modified on 03/16/17
    3812576257300004
    [42] Apteryx
    Taxon name last Modified on 03/16/17
    4182540253600004
    [43] Eudromia
    Taxon name last Modified on 03/16/17
    2212741273300004
    [44] Chauna
    Taxon name last Modified on 03/16/17
    1492809280500004
    [45] Anseranas
    Taxon name last Modified on 03/16/17
    1652791278900004
    [46] Anser
    Taxon name last Modified on 03/16/17
    1522806280200004
    [47] Anas
    Taxon name last Modified on 03/16/17
    1462813280800004
    [48] Megapodius
    Taxon name last Modified on 03/16/17
    1412815281300004
    [49] Ortalis
    Taxon name last Modified on 03/16/17
    1382818281600004
    [50] Meleagris
    Taxon name last Modified on 03/16/17
    1402816281400004
    [51] Dendragapus
    Taxon name last Modified on 03/16/17
    1432814281100004
    [52] Lophortyx
    Taxon name last Modified on 03/16/17
    1432814281100004
    [53] Francolinus
    Taxon name last Modified on 03/16/17
    1432813281100004
    [54] Alectoris
    Taxon name last Modified on 03/16/17
    1472809280700004
    [55] Gallus
    Taxon name last Modified on 03/16/17
    1432813281100004
    [56] Numida
    Taxon name last Modified on 12/05/13
    1482809280600004
    [57] Sylviornis
    Taxon name last Modified on 12/05/13
    270025525400004
    [58] Diatryma
    Taxon name last Modified on 12/05/13
    281214314200004
    [59] Dromornithidae
    Taxon name last Modified on 03/16/17
    17591201119500004
    [60] Diomedea
    Taxon name last Modified on 03/16/17
    1492813280500004
    [61] Puffinus
    Taxon name last Modified on 03/16/17
    1522804280200004
    [62] Pachyptila
    Taxon name last Modified on 03/16/17
    1522804280200004
    [63] Oceanites
    Taxon name last Modified on 03/16/17
    1562800279800004
    [64] Pelecanoides
    Taxon name last Modified on 03/16/17
    1722783278200004
    [65] Spheniscus
    Taxon name last Modified on 03/16/17
    2592697269500004
    [66] Gavia
    Taxon name last Modified on 03/16/17
    1612793279300004
    [67] Podiceps
    Taxon name last Modified on 03/16/17
    1722785278200004
    [68] Fregata
    Taxon name last Modified on 03/16/17
    1672788278700004
    [69] Anhinga
    Taxon name last Modified on 03/16/17
    1622792279200004
    [70] Phalacrocorax
    Taxon name last Modified on 03/16/17
    1622793279200004
    [71] Sula
    Taxon name last Modified on 12/05/13
    1692785278500004
    [72] Pelecanus
    Taxon name last Modified on 03/16/17
    1662788278800004
    [73] Phaethon
    Taxon name last Modified on 03/16/17
    1662789278800004
    [74] Ardea
    Taxon name last Modified on 03/16/17
    1562799279800004
    [75] Nycticorax
    Taxon name last Modified on 03/16/17
    1492806280500004
    [76] Tigrisoma
    Taxon name last Modified on 03/16/17
    1522803280200004
    [77] Botaurus
    Taxon name last Modified on 03/16/17
    1582797279600004
    [78] Cochlearius
    Taxon name last Modified on 12/05/13
    1522804280200004
    [79] Ajaia
    Taxon name last Modified on 03/16/17
    1512804280300004
    [80] Scopus
    Taxon name last Modified on 03/16/17
    1532801280100004
    [81] Plegadis
    Taxon name last Modified on 03/16/17
    1602795279400004
    [82] Ciconia
    Taxon name last Modified on 03/16/17
    1432812281100004
    [83] Phoenicopterus
    Taxon name last Modified on 03/16/17
    1582800279600004
    [84] Balaeniceps
    Taxon name last Modified on 03/16/17
    1762779277800004
    [85] Accipiter
    Taxon name last Modified on 03/16/17
    1552800279900004
    [86] Gyps
    Taxon name last Modified on 03/16/17
    1712783278300004
    [87] Pandion
    Taxon name last Modified on 03/16/17
    1492805280500004
    [88] Falco
    Taxon name last Modified on 03/16/17
    1462808280800004
    [89] Polyborus
    Taxon name last Modified on 03/16/17
    1422812281200004
    [90] Sagittarius
    Taxon name last Modified on 03/16/17
    1532801280100004
    [91] Catharthes
    Taxon name last Modified on 12/05/13
    1512803280300004
    [92] Turnix
    Taxon name last Modified on 03/16/17
    1812773277300004
    [93] Mesitornis
    Taxon name last Modified on 03/16/17
    1962758275800004
    [94] Afrotis
    Taxon name last Modified on 03/16/17
    1632793279100004
    [95] Cariama
    Taxon name last Modified on 03/16/17
    1552799279900004
    [96] Eurypyga
    Taxon name last Modified on 03/16/17
    1412813281300004
    [97] Rhynochetos
    Taxon name last Modified on 03/16/17
    1642790279000004
    [98] Aptornis
    Taxon name last Modified on 12/05/13
    9631995199100004
    [99] Psophia
    Taxon name last Modified on 03/16/17
    1522802280200004
    [100] Aramus
    Taxon name last Modified on 03/16/17
    1532802280100004
    [101] Grus
    Taxon name last Modified on 03/16/17
    1492809280500004
    [102] Heliornis
    Taxon name last Modified on 03/16/17
    2052754274900004
    [103] Porphyrula
    Taxon name last Modified on 03/16/17
    1482810280600004
    [104] Pedionomus
    Taxon name last Modified on 03/16/17
    2052750274900004
    [105] Jacana
    Taxon name last Modified on 03/16/17
    1412814281300004
    [106] Rostratula
    Taxon name last Modified on 03/16/17
    1492807280500004
    [107] Dromas
    Taxon name last Modified on 03/16/17
    1812774277300004
    [108] Haematopus
    Taxon name last Modified on 03/16/17
    1612794279300004
    [109] Ibidorhyncha
    Taxon name last Modified on 03/16/17
    1982757275600004
    [110] Himantopus
    Taxon name last Modified on 03/16/17
    1732783278100004
    [111] Cladorhynchus
    Taxon name last Modified on 03/16/17
    2242731273000004
    [112] Burhinus
    Taxon name last Modified on 03/16/17
    1742781278000004
    [113] Glareola
    Taxon name last Modified on 03/16/17
    1592800279500004
    [114] Cursorius
    Taxon name last Modified on 03/16/17
    1792776277500004
    [115] Pluvialis
    Taxon name last Modified on 03/16/17
    1692794278500004
    [116] Chionis
    Taxon name last Modified on 03/16/17
    1572798279700004
    [117] Heteroscelus
    Taxon name last Modified on 03/16/17
    1472815280700004
    [118] Phalaropus
    Taxon name last Modified on 03/16/17
    1462810280800004
    [119] Thinocorus
    Taxon name last Modified on 03/16/17
    1592796279500004
    [120] Uria
    Taxon name last Modified on 03/16/17
    1732785278100004
    [121] Stercorarius
    Taxon name last Modified on 03/16/17
    1552801279900004
    [122] Rissa
    Taxon name last Modified on 03/16/17
    1682794278600004
    [123] Chlidonias
    Taxon name last Modified on 03/16/17
    1512809280300004
    [124] Rynchops
    Taxon name last Modified on 03/16/17
    1662789278800004
    [125] Pterocles
    Taxon name last Modified on 03/16/17
    1562800279800004
    [126] Columba
    Taxon name last Modified on 03/16/17
    1472809280700004
    [127] Zenaida
    Taxon name last Modified on 12/05/13
    1492806280500004
    [128] Geopelia
    Taxon name last Modified on 03/16/17
    1512804280300004
    [129] Didunculus
    Taxon name last Modified on 03/16/17
    1742781278000004
    [130] Goura
    Taxon name last Modified on 03/16/17
    1502806280400004
    [131] Ducula
    Taxon name last Modified on 03/16/17
    1532802280100004
    [132] Raphus
    Taxon name last Modified on 03/16/17
    7252229222900004
    [133] Pezophaps
    Taxon name last Modified on 03/16/17
    7182236223600004
    [134] Trichoglossus
    Taxon name last Modified on 03/16/17
    1812774277300004
    [135] Cacatua
    Taxon name last Modified on 03/16/17
    1892766276500004
    [136] Amazona
    Taxon name last Modified on 03/16/17
    1802775277400004
    [137] Melopsittacus
    Taxon name last Modified on 03/16/17
    1872768276700004
    [138] Cuculus
    Taxon name last Modified on 03/16/17
    1512803280300004
    [139] Coccyzus
    Taxon name last Modified on 03/16/17
    1572797279700004
    [140] Crotophaga
    Taxon name last Modified on 03/16/17
    1472807280700004
    [141] Geococcyx
    Taxon name last Modified on 03/16/17
    1602794279400004
    [142] Centropus
    Taxon name last Modified on 03/16/17
    1582796279600004
    [143] Corythaixoides
    Taxon name last Modified on 03/16/17
    1532801280100004
    [144] Opisthocomus
    Taxon name last Modified on 03/16/17
    1472807280700004
    [145] Tyto
    Taxon name last Modified on 03/16/17
    1502804280400004
    [146] Phodilus
    Taxon name last Modified on 03/16/17
    2072747274700004
    [147] Otus
    Taxon name last Modified on 03/16/17
    1492805280500004
    [148] Strix
    Taxon name last Modified on 03/16/17
    1532801280100004
    [149] Steatornis
    Taxon name last Modified on 03/16/17
    1632791279100004
    [150] Podargus
    Taxon name last Modified on 03/16/17
    1862768276800004
    [151] Nyctibius
    Taxon name last Modified on 03/16/17
    1622792279200004
    [152] Aegotheles
    Taxon name last Modified on 03/16/17
    2032751275100004
    [153] Caprimulgus
    Taxon name last Modified on 03/16/17
    1502804280400004
    [154] Hemiprocne
    Taxon name last Modified on 03/16/17
    1862768276800004
    [155] Cypseloides
    Taxon name last Modified on 03/16/17
    1902767276400004
    [156] Apus
    Taxon name last Modified on 03/16/17
    1722782278200004
    [157] Glaucis
    Taxon name last Modified on 03/16/17
    1932762276100004
    [158] Colius
    Taxon name last Modified on 03/16/17
    1642790279000004
    [159] Pharomachrus
    Taxon name last Modified on 03/16/17
    1612793279300004
    [160] Trogon
    Taxon name last Modified on 03/16/17
    1562798279800004
    [161] Alcedo
    Taxon name last Modified on 12/05/13
    1682786278600004
    [162] Halcyon
    Taxon name last Modified on 03/16/17
    1772777277700004
    [163] Todus
    Taxon name last Modified on 03/16/17
    1732781278100004
    [164] Momotus
    Taxon name last Modified on 03/16/17
    1712783278300004
    [165] Merops
    Taxon name last Modified on 03/16/17
    1562798279800004
    [166] Coracias
    Taxon name last Modified on 03/16/17
    1532801280100004
    [167] Brachypteracias
    Taxon name last Modified on 03/16/17
    2322722272200004
    [168] Leptosomus
    Taxon name last Modified on 12/05/13
    1932761276100004
    [169] Upupa
    Taxon name last Modified on 03/16/17
    1842770277000004
    [170] Phoeniculus
    Taxon name last Modified on 03/16/17
    2012753275300004
    [171] Tockus
    Taxon name last Modified on 03/16/17
    1642792279000004
    [172] Galbula
    Taxon name last Modified on 03/16/17
    1752782277900004
    [173] Monasa
    Taxon name last Modified on 03/16/17
    1572797279700004
    [174] Megalaima
    Taxon name last Modified on 03/16/17
    1812774277300004
    [175] Semornis
    Taxon name last Modified on 12/05/13
    1812775277300004
    [176] Indicator
    Taxon name last Modified on 03/16/17
    1832771277100004
    [177] Aulocorhynchus
    Taxon name last Modified on 12/05/13
    1852770276900004
    [178] Ramphastos
    Taxon name last Modified on 03/16/17
    1732782278100004
    [179] Jynx
    Taxon name last Modified on 03/16/17
    1942760276000004
    [180] Picoides
    Taxon name last Modified on 03/16/17
    1832776277100004
    [181] Menura
    Taxon name last Modified on 03/16/17
    1992756275500004
    [182] Pitta
    Taxon name last Modified on 03/16/17
    1832772277100004
    [183] Pitangus
    Taxon name last Modified on 03/16/17
    1592796279500004
    [184] Ptilonorhynchus
    Taxon name last Modified on 03/16/17
    1622793279200004
    [185] Aphelocoma
    Taxon name last Modified on 03/16/17
    1562799279800004
    [186] Bombycilla
    Taxon name last Modified on 03/16/17
    1632792279100004
    [187] Parus
    Taxon name last Modified on 03/16/17
    1782777277600004
    [188] Passer
    Taxon name last Modified on 03/16/17
    1732782278100004


    Project downloads

    type number of downloads Individual items downloaded (where applicable)
    Total downloads from project1346
    Document downloads176Matrix without character names (49 downloads); Livezey Zusi codings (60 downloads); Read Me (32 downloads); Standard Assumptions (19 downloads); Standard Ordering (16 downloads);
    Matrix downloads295Morphology Matrix (295 downloads);
    Project downloads872
    Media downloads3M437548 (1 download); M437549 (1 download); M88756 (1 download);