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Zhonghe Zhou, Fu-cheng Zhang (2001)
Two new ornithurine birds from the Early Cretaceous of western Liaoning, ChinaChinese Science Bulletin, 46
J. Baumel (1988)
Functional morphology of the tail apparatus of the pigeon (Columba livia).Advances in anatomy, embryology, and cell biology, 110
length: (0) shorter than humerus, (1) as long as or longer than the humerus
(2002)
Locomotor evolution on the line to modern birds. In: Mesozoic Birds: Above the Heads of Dinosaurs (eds Chiappe LM
(1993)
Osteologia. In: Handbook of Avian Anatomy: Nomina Anatomica Avium, 2nd edn (eds Baumel JJ, King AS
Zhonghe Zhou, J. Clarke, Fu-cheng Zhang (2002)
Archaeoraptor's better halfNature, 420
) fused, (1) unfused
L. Martin, D. Steadman, P. Rich (1983)
Perspectives in ornithology: The origin and early radiation of birds
Preacetabular ilium extends anterior to first sacral vertebrae: (0) no free ribs overlapped, (1) one or more ribs overlapped
Femur, fossa for insertion of lig
Metatarsal III, trochlea in plantar view, proximal extent of lateral and medial edges of trochlea: (0) absent, trochlear edges approximately equal in proximal extent
Semilunate carpal and metacarpals: (0) no fusion, (1) incomplete proximal fusion
Distal end of pubis: (0) expanded, flared; (1) straight, subequal, in proportion with rest of pubis
) subcylindrical to subtriangular; (1) strongly dorsoventrally compressed, flat caudal surface
M. Evans, Adrian Thomas (1992)
The aerodynamic and mechanical effects of elongated tails in the scarlet-tufted malachite sunbird: measuring the cost of a handicapAnimal Behaviour, 43
metatarsal II shorter than metatarsal IV, but reaching distally further than base of metatarsal IV trochlea
Preacetabular ilium: (0) approach on midline, open, or cartilaginous connection, (1) co-ossified, dorsal closure of 'iliosynsacral canals
) II approximately the same size as III and/or IV, (1) II wider than III and/or IV
Femur: (0) ectocondylar tubercle and lateral condyle separated by deep notch, (1) ectocondylar tubercle and lateral condyle form single trochlear surface
G. Dyke, R. Dortangs, J. Jagt, E. Mulder, A. Schulp, L. Chiappe (2002)
Europe's last Mesozoic birdNaturwissenschaften, 89
) straight, (1) dorsoventrally curved
) approximately equal, (2) lateral condyle wider (Ordered)
M. Norell, J. Clarke (2001)
Fossil that fills a critical gap in avian evolutionNature, 409
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The morphology and taxonomy of Ichthyornis Marsh and the phylogenetic relationships of basal Ornithurae
Adrian Thomas (1993)
On the aerodynamics of birds’ tailsPhilosophical transactions - Royal Society. Mathematical, physical and engineering sciences, 340
) pit-shaped scapular cotyla developed on the coracoid, and coracoidal tubercle developed on the scapula ('ball and socket' articulation); (1) scapular articular surface of coracoid convex
L. Hou, L. Martin, Zhonghe Zhou, A. Feduccia (1996)
Early Adaptive Radiation of Birds: Evidence from Fossils from Northeastern ChinaScience, 274
S. Gatesy, K. Dial (1996)
FROM FROND TO FAN: ARCHAEOPTERYX AND THE EVOLUTION OF SHORT‐TAILED BIRDSEvolution, 50
) length of long axis of condyle less than the same measure of the dorsal condyle
Intermetacarpal process or tubercle: (0) absent, (1) present as scar
D. Steadman (2003)
Mesozoic Birds: Above the Heads of Dinosaurs, 120
) absent; (1) present, developed as a flat scar or as a scar-impressed fossa
Postacetabular ilium, ventral surface
Hou Hou, Zhang Zhang (1993)
A new fossil bird from the Lower Cretaceous of LiaoningVert Pal Asiat, 31
posterior demarcation of the obturator foramen: (0) absent; (1) present, developed as a small flange or raised scar contacting/fused with pubis and demarcating the obturator foramen distally
Ossified uncinate processes: (0) absent, (1) present and unfused to ribs
Ulnare: (0) absent, (1) present
Angle between coracoid and scapula at glenoid: (0) more than 90 degrees, (1) 90 degrees or less
Zhonghe Zhou (2004)
The origin and early evolution of birds: discoveries, disputes, and perspectives from fossil evidenceNaturwissenschaften, 91
Huaiyu He, Xiaolin Wang, Zhonghe Zhou, F. Wang, A. Boven, G. Shi, R. Zhu (2004)
Timing of the Jiufotang Formation (Jehol Group) in Liaoning, northeastern China, and its implicationsGeophysical Research Letters, 31
Gatesy Gatesy, Dial Dial (1993)
Tail muscle activity patterns in walking and flying pidgeonsColumba livia. J Exp Biol, 176
J. Clarke, M. Norell (2004)
New Avialan Remains and a Review of the Known Avifauna from the Late Cretaceous Nemegt Formation of Mongolia
Chiappe Chiappe (1995b)
The phylogenetic position of the Cretaceous birds of Argentina: Enantiornithes and Patagopteryx deferrariisiCour Forsch Senckenberg, 181
) absent, (1) a tubercle, (2) an elongate process (Ordered)
J. Hutchinson (2001)
The evolution of femoral osteology and soft tissues on the line to extant birds (Neornithes)Zoological Journal of the Linnean Society, 131
Ischium: (0) forked (dorsal process present); (1) straight, no dorsal process
Louise Rowe, M. Evans, K. Buchanan (2001)
The function and evolution of the tail streamer in hirundinesBehavioral Ecology, 12
) a groove open distally and bounded medially and laterally by narrow condyles, (1) closed distally by expansion of both condyles
A. Barbosa, A. Møller (1999)
Aerodynamic costs of long tails in male barn swallows Hirundo rustica and the evolution of sexual size dimorphismBehavioral Ecology, 10
D. Swofford, D. Swofford, D. Swofford (2002)
PAUP*: Phylogenetic analysis using parsimony (*and other methods), Version 4.0b10
J. Rayner (1988)
Form and Function in Avian Flight
) no indication of origin as a scar, a pit, or a tubercle; (1) indication as a pit-shaped scar or as a variably projected scar-bearing tubercle or facet
Intermetacarpal space (between metacarpals II and III) (0) reaches proximally as far as the distal end of metacarpal I, (1) terminates distal to end of metacarpal I
M. Norell, P. Makovicky, Mongolyn Akademi. (1997)
Important features of the dromaeosaur skeleton : information from a new specimen. American Museum novitates ; no. 3215
E. Kurochkin, G. Dyke, A. Karhu (2002)
A New Presbyornithid Bird (Aves, Anseriformes) from the Late Cretaceous of Southern Mongolia
Calcaneum and astragalus: (0) unfused to each other or tibia in adult
Laterally projected fibular trochlea: (0) absent; (1) present, developed as small notch
ventral margin of apophysis: (0) curved, angling; (1) with a truncate or squared base
Zhonghe Zhou, P. Barrett, J. Hilton (2003)
An exceptionally preserved Lower Cretaceous ecosystemNature, 421
Tibia/tarsal formed condyles, extensor canal: (0) absent, (1) an emarginate groove
metacarpals II and III, articular surfaces for digits: (0) metacarpal II subequal or surpasses metacarpal III in distal extent
Ulnare: (0), 'heart-shaped', little differentiation into short dorsal and ventral rami, (1) V-shaped, welldeveloped dorsal and ventral rami
Zhang Zhang, Zhou Zhou (2004)
Leg feathers in an Early Cretaceous birdNature, 431
) roughly hourglassshaped proximally, at least moderately expanded
dorsal surface (= posterior surface of basal maniraptoran theropods): (0) strongly concave, (1) flat to convex
Pubis: (0) suboval in cross section
Hou Hou, Martin Martin, Zhou Zhou, Feduccia Feduccia (1996)
Early adaptation of birds‐evidence from fossils from Northeastern ChinaScience, 27
Marsh Marsh (1872b)
Notice of a new and remarkable fossil birdAm J Sci, 3rd Ser, 4
O. Marsh (1872)
Preliminary description of Hesperornis regalis, with notice of four other new species of Cretaceous birdsAmerican Journal of Science, s3-3
Alvarenga Alvarenga, Bonaparte Bonaparte (1992)
A new flightless bird from the Cretaceous of PatagoniaNat Hist Mus Los Angeles, Sci Series, 36
J. Clarke (2004)
MORPHOLOGY, PHYLOGENETIC TAXONOMY, AND SYSTEMATICS OF ICHTHYORNIS AND APATORNIS (AVIALAE: ORNITHURAE)
proximal vascular foramen (foramina): (0) absent, (1) one, between metatarsals III and IV
L. Chiappe (1996)
Late Cretaceous birds of southern South America: anatomy and systematics of Enantiornithes and Patagopteryx deferrariisi, 30
L. Hou, L. Chiappe, Fu-cheng Zhang, C. Chuong (2003)
New Early Cretaceous fossil from China documents a novel trophic specialization for Mesozoic birdsNaturwissenschaften, 91
(1983)
) distal tarsals fuse to metatarsals, and metatarsals fuse to each other proximally and distally
S. Olson (2000)
Anatomy and Systematics of the Confuciusornithidae (Theropoda: Aves) from the Late Mesozoic of Northeastern China, 117
S. Gatesy, K. Dial (1993)
TAIL MUSCLE ACTIVITY PATTERNS IN WALKING AND FLYING PIGEONS (COLUMBA LIVIA)The Journal of Experimental Biology, 176
distal plantar surface, fossa for metatarsal I (fossa metatarsi I; Baumel and Witmer, 1993): (0) absent, (1) shallow notch, (2) conspicuous ovoid fossa (Ordered)
J. Cracraft (1986)
The origin and early diversification of birdsPaleobiology, 12
cnemial crest(s): (0) lateral crest only, (1) lateral and anterior crests developed
Fu-cheng Zhang, Zhonghe Zhou (2004)
Palaeontology: Leg feathers in an Early Cretaceous birdNature, 431
Zhonghe Zhou, J. Clarke, Fu-cheng Zhang, O. Wings (2004)
Gastroliths in Yanornis: an indication of the earliest radical diet-switching and gizzard plasticity in the lineage leading to living birds?Naturwissenschaften, 91
Interclavicular angle (clavicles elongate): (0) greater than, or equal
weakly defined, 'strap-like
) proximally in plane with II and IV, (1) proximally displaced plantarly, relative to metatarsals II and IV
(1993)
dorsal ramus (crus brevis), (1) same length as dorsal ramus, (2) longer than dorsal ramus. 141 Semilunate carpal and metacarpals
Carina or midline ridge: (0) restricted to posterior half of sternum, (1) approaches anterior limit of sternum
L. Chiappe (1995)
The first 85 million years of avian evolutionNature, 378
Clarke Clarke, Norell Norell, Zhou Zhou, Zhang Zhang (2002)
An ornithurine from the Early Cretaceous of ChinaJ Vert Paleontol, 22
K. Bremer (1988)
THE LIMITS OF AMINO ACID SEQUENCE DATA IN ANGIOSPERM PHYLOGENETIC RECONSTRUCTIONEvolution, 42
J. Clarke, C. Tambussi, J. Noriega, G. Erickson, R. Ketcham (2005)
Definitive fossil evidence for the extant avian radiation in the CretaceousNature, 433
(2006)
The Authors Journal compilation
Zhonghe Zhou, Fu-cheng Zhang (2003)
Anatomy of the primitive bird Sapeornis chaoyangensis from the Early Cretaceous of Liaoning, ChinaCanadian Journal of Earth Sciences, 40
J. Clarke, M. Norell (2002)
The Morphology and Phylogenetic Position of Apsaravis ukhaana from the Late Cretaceous of Mongolia
Laterally projected process on ischiadic peduncle (antitrochanter): (0) directly posterior to acetabulum
L. Hou, L. Martin, Zhonghe Zhou, A. Feduccia, Fu-cheng Zhang (1999)
A diapsid skull in a new species of the primitive bird ConfuciusornisNature, 399
) contacting, variably coossified into symphysis
) same length as dorsal ramus
Manual digit II, phalanges: (0) length of phalanx II-1 less than or equal to that of II-2
(1996)
The discovery of a Jurassic carinate bird in China
(1985)
Lower Cretaceous birds from Mongolia and their evolutionary significance
J. Gauthier (1986)
Saurischian monophyly and the origin of birds, 8
L. Chiappe (1991)
Cretaceous avian remains from Patagonia shed new light on the early radiation of birdsAlcheringa, 15
A. Keast (1996)
WING SHAPE IN INSECTIVOROUS PASSERINES INHABITING NEW GUINEA AND AUSTRALIAN RAIN FORESTS AND EUCALYPT FOREST/EUCALYPT WOODLANDSThe Auk, 113
Metacarpal I, distal articulation with phalanx I: (0) ginglymoid, (1) shelf
dorsal trochlear surface, extent along posterior margin: (0) less than transverse measure of dorsal trochlear surface, (1) approximately equal in extent
Aves): (0) absent; (1) present, on approximately the center of the proximodorsal surface of metatarsal II
A. Balmford, I. Jones, Adrian Thomas (1993)
On avian asymmetry: evidence of natural selection for symmetrical tails and wings in birdsProceedings of the Royal Society of London. Series B: Biological Sciences, 252
Ernst Haeckel (1866)
Generelle Morphologie der Organismen: Allgemeine Grundzüge der organischen Formen-Wissenschaft, mechanisch begründet durch die von Charles Darwin reformierte Descendenz-Theorie. Band 1: Allgemeine Anatomie. Band 2: Allgemeine Entwicklungsgeschichte
) developed distally, (1) developed on anterior surface of humerus
Chang Meemann (2003)
The Jehol Biota: the emergence of feathered dinosaurs, beaked birds and flowering plants
Ernst Haeckel
Generelle Morphologie der Organismen, 1
) present. 89 Coracoid: (0) height approximately equal mediolateral dimension; (1) height more than twice width
humerotricipitalis groove: (0) absent, (1) present as a ventral depression contiguous with the olecranon fossa
R. Barsbold, P. Currie, N. Myhrvold, H. Osmólska, K. Tsogtbaatar, M. Watabe (2000)
A pygostyle from a non-avian theropodNature, 403
(2001)
) present, developed as a slightly projected tubercle or flange; (1) hypertrophied, 'shelf-like' conformation (in combination with development of the trochanteric shelf; see Hutchinson
area of the foramen n. supracoracoideus (when developed): (0) strongly depressed
Hou Hou, Liu Liu (1984)
A new fossil bird from the Lower Cretaceous of Gansu and early evolution of birdsSci Sinica (B), 27
) absent, (1) developed as a slightly raised scar
relative mediolateral width: (0) metatarsal IV approximately the same width as metatarsals II and III, (1) metatarsal IV narrower than MII and MIII
) widely separated mediolaterally, (1) adjacent, (2) crossed on midline
Norell Norell, Mackovicky Mackovicky (1997)
Important features of the dromaeosaur skeleton: information from a new specimenAm Mus Novit, 3215
pit-shaped scar/fossa for muscular attachment on anterodistal, distal or posterodistal surface of crest: (0) absent
Sternum, posterior margin, distinct posteriorly projected medial and/or lateral processes: (0) absent new clade)
Zhang Zhang, Zhou Zhou (2000)
A primitive enantiornithine bird and the origin of feathersScience, 290
Lianhai Hou, Zhicheng Liu (1984)
A NEW FOSSIL BIRD FROM LOWER CRETACEOUS OF GANSU AND EARLY EVOLUTION OF BIRDSScience in China Series B-Chemistry, Biological, Agricultural, Medical & Earth Sciences, 27
) little or no anterior projection, (1) developed as an anterior projection relative to shaft surface in ventral view
distal extent of metatarsal II relative to metatarsal IV: (0) approximately equal in distal extent
O. Marsh
Notice of a new and remarkable fossil birdJournal of Natural History, 11
Postacetabular ilium: (0) dorsoventrally orientated, (1) mediolaterally orientated
Zhonghe Zhou, Fu-cheng Zhang (2001)
Largest bird from the Early Cretaceous and its implications for the earliest avian ecological diversificationNaturwissenschaften, 89
(2004)
New avialan remains from the Late Cretaceous of Mongolia and a review of the known avifauna of the Nemegt Formation
acrocoracoid: (0) straight, (1) hooked medially
Ulna, cotylae: (0) dorsoventrally adjacent, (1) widely separated by a deep groove
) not subparallel, pubis directed nearly directly ventrally; (1) subparallel, pubis posteriorly directed
) dorsal to, or at approximately same level as
Fitzpatrick (1999)
Tail length in birds in relation to tail shape, general flight ecology and sexual selectionJournal of Evolutionary Biology, 12
dorsal (omal) tip: (0) flat or blunt tip
number of processes for articulation with the sternal ribs: (0) three, (1) four, (2) five, (3) six, (4) seven or more (Ordered)
Sternum: raised, paired intermuscular ridges
K. Padian, L. Chiappe (1998)
The origin and early evolution of birdsBiological Reviews, 73
J. Clarke, L. Chiappe (2001)
A New Carinate Bird from the Late Cretaceous of Patagonia (Argentina)
) well developed, globose
) wider or approximately the same width as proximal dorsoventral shaft width
K. Buchanan, M. Evans (2000)
The effect of tail streamer length on aerodynamic performance in the barn swallowBehavioral Ecology, 11
S. Gatesy, K. Dial (1996)
LOCOMOTOR MODULES AND THE EVOLUTION OF AVIAN FLIGHTEvolution, 50
Distal caudals: (0) unfused, (1) fused
O. Savile (1957)
ADAPTIVE EVOLUTION IN THE AVIAN WINGEvolution, 11
) smooth, (1) with muscle impression along most of surface
In studies of the evolution of avian flight there has been a singular preoccupation with unravelling its origin. By contrast, the complex changes in morphology that occurred between the earliest form of avian flapping flight and the emergence of the flight capabilities of extant birds remain comparatively little explored. Any such work has been limited by a comparative paucity of fossils illuminating bird evolution near the origin of the clade of extant (i.e. ‘modern’) birds (Aves). Here we recognize three species from the Early Cretaceous of China as comprising a new lineage of basal ornithurine birds. Ornithurae is a clade that includes, approximately, comparatively close relatives of crown clade Aves (extant birds) and that crown clade. The morphology of the best‐preserved specimen from this newly recognized Asian diversity, the holotype specimen of Yixianornis grabaui Zhou and Zhang 2001, complete with finely preserved wing and tail feather impressions, is used to illustrate the new insights offered by recognition of this lineage. Hypotheses of avian morphological evolution and specifically proposed patterns of change in different avian locomotor modules after the origin of flight are impacted by recognition of the new lineage. The complete articulated holotype specimen of Yixianornis grabaui, from the Early Cretaceous Jiufotang Formation of Liaoning Province, in north‐eastern China, arguably the best‐preserved basal ornithurine specimen yet discovered, provides the earliest evidence consistent with the presence of extant avian tail feather fanning.
Journal of Anatomy – Wiley
Published: Mar 1, 2006
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