A second enantiornithean (Aves: Ornithothoraces) wing from the Early Cretaceous Xiagou Formation near Changma, Gansu Province, People's Republic of China

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    A second enantiornithean (Aves: Ornithothoraces)wing from the Early Cretaceous Xiagou Formationnear Changma, Gansu Province, Peoples Republicof China

    Jerald D. Harris, Matthew C. Lamanna, Hai-lu You, Shu-an Ji, and Qiang Ji

    Abstract: A new specimen of an enantiornithean bird from the Lower Cretaceous Xiagou Formation of Gansu Province,northwestern China, consists of an articulated distal left humerus, ulna, radius, carpus, and manus. The specimen mayrepresent a primitive enantiornithean because it lacks a longitudinal sulcus on the radius, has incompletely fused alularand major metacarpals, and possibly retains a remnant of a second phalanx on the minor digit. It differs from all otherknown enantiornitheans, and exhibits possible autapomorphies, including peculiar, flat humeral epicondyles, a pair ofeminences on the distal minor metacarpal, and an enormous flexor tuberculum on the alular ungual. The specimenprobably pertains to the same taxon as a previously described enantiornithean arm from Changma; the incompletenessof the taxon precludes erecting a new name, but it provides new information concerning enantiornithean diversity in theEarly Cretaceous of central Asia.

    Rsum : Un nouveau spcimen dun oiseau enantiornithien de la Formation de Xiagou du Crtac infrieur de laprovince de Gansu, du nord-ouest de la Chine, consiste en un humrus distal gauche articul, un cubitus, un radius, uncarpe et une paume. Labsence dun sillon longitudinal sur le radius, la fusion incomplte de lalulaire et des mtacarpiensmajeurs, ainsi que la prsence possible dune deuxime phalange vestigiale sur le doigt mineur pourraient indiquer quilsagit dun enantiornithien primitif. Le spcimen se distingue de tous les autres enantiornithiens connus et prsentepossiblement des autapomorphies dont de singuliers picondyles aplatis de lhumrus, une paire dminences sur lemtacarpien mineur distal et un norme tubercule flchisseur sur lungual alulaire. Ce spcimen est probablementassoci au mme taxon quun bras enantiornithien provenant de Changma dcrit prcdemment. tant donn la natureincomplte du taxon, il est impossible de proposer un nouveau nom, mais le spcimen fournit toutefois des nouvellesdonnes sur la diversit des oiseaux enantiornithiens durant le Crtac prcoce en Asie centrale.

    [Traduit par la Rdaction] Harris et al. 554


    Taxa pertaining to the avian clade Enantiornithes are widelyrecognized as the most common and diverse birds of theCretaceous Period. Though the first discovered fossils nowattributed to the clade are from Upper Cretaceous sediments(Brodkorb 1976; Elzanowski 1976, 1977; Walker 1981), mostof the best known enantiornitheans have been recovered fromLower Cretaceous sediments, particularly in Spain (Sanz etal. 2002) and Jehol Biota-bearing deposits in northeasternChina (Gong et al. 2004; Hou et al. 2004; Zhang et al. 2004;Zhou and Hou 2002). Historically, however, Early Cretaceousbirds in China were initially represented by the fragmentary

    non-enantiornithean ornithothoracean Gansus yumenensis fromthe Xiagou Formation in western Gansu Province, north-western China (Hou and Liu 1984). Following its description,a nearly two-decade long hiatus followed, in which no newbirds were discovered in Gansu.

    In 2004, a collaborative expedition led by researchers fromthe Chinese Academy of Geological Sciences and CarnegieMuseum of Natural History revisited these Xiagou Formationoutcrops (Fig. 1) and recovered numerous partial to nearlycomplete avian skeletons, several preserving feather and soft-tissue impressions. Of these, two enantiornithean specimenshave already been briefly described (OConnor et al. 2004;Lamanna et al. 2005; You et al. 2005). Here a third speci-

    Can. J. Earth Sci. 43: 547554 (2006) doi:10.1139/E06-007 2006 NRC Canada

    Received 17 October 2005. Accepted 23 January 2006. Published on the NRC Research Press Web site at http://cjes.nrc.ca on18 May 2006.

    Paper handled by Associate Editor H.-D. Sues.

    J.D. Harris.1 Science Department, Dixie State College, 225 South 700 East, St. George, UT 84770, USA.M.C. Lamanna. Section of Vertebrate Paleontology, Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, PA15213-4080, USA.H.-l. You, S.-a. Ji, Q. Ji. Institute of Geology, Chinese Academy of Geological Sciences, 26 Baiwanzhuang Road, Beijing 100037,P.R. China.

    1Corresponding author (e-mail jharris@dixie.edu).

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    men, an incomplete, articulated forelimb of a finch-sizedenantiornithean, is described.

    The specimen is clearly avian because it possesses a carpo-metacarpus in which the semilunate carpal is fused to atleast the major and minor metacarpals (Chiappe 2002). Itdemonstrably pertains to the Ornithothoraces because its alulardigit does not surpass the major metacarpal in length (Chiappe2002). The specimen can be assigned to the Enantiornithesbecause its minor metacarpal extends further distally thanthe major metacarpal. The radius appears to lack the sulcuson its interosseous surface that may be autapomorphic of theEuenantiornithes (Chiappe and Walker 2002). The specimenalso lacks diagnostic ornithuromorph characters, such as awide interosseous space and a carpometacarpus with distallyfused major and minor metacarpals. Three-dimensional pres-ervation allows for especially detailed morphological obser-vations.

    TerminologyBecause the specimen described herein pertains to a bird,

    osteological terminology used herein follows Baumel andWitmer (1993). The term enantiornithean, rather thanenantiornithine, is used herein as an informal shorthandfor a member of the clade Enantiornithes because the latterimplies the existence of, and membership in, a cladeEnantiornithinae despite the fact that no such clade hasever been recognized. Similarly, ornithothoracean is usedinstead of ornithothoracine because there is no cladeOrnithothoracinae, only Ornithothoraces. The same logicapplies to ornithuran versus ornithurine for Ornithurae,as well as neornithean versus neornithine for Neornithesand galloanseran versus galloanserine for Galloanserae,although the latter clade is not discussed herein. This patternconforms to prevalent usage for other avian and nonaviantheropod clade names with similar suffixes (e.g., avianrather than avine for Aves, avialan rather than avialinefor Avialae, and tetanuran rather than tetanurine forTetanurae) and brings paleornithological terminology intocongruence with the nomenclature of other organisms (e.g.,gnetalean for Gnetales, aranean for Araneae, schizaceanfor Schizaceae, etc.).

    AbbreviationsCAGS-IG, Chinese Academy of Geological Sciences,

    Institute of Geology, Beijing, China.

    Systematic Paleontology

    Aves Linneus, 1758 (Avialae sensu Gauthier 1986)Pygostylia Chatterjee, 1997 (sensu Chiappe 2002)Ornithothoraces Chiappe, 1995Enantiornithes Walker, 1981Taxon indet.(Figs. 24)

    SPECIMEN: CAGS-IG-04-CM-023 (hereinafter CM-023 for thesake of brevity), an incomplete, articulated left thoracic limb,consists of a distal humerus and complete ulna, radius, carpus,carpometacarpus, and manual digits.

    LOCALITY: Near the town of Changma in the Changma Basinof northwestern Gansu Province, China (Fig. 1).

    HORIZON: Lower Cretaceous (?AptianAlbian) Xiagou Forma-tion, middle unit of the Xinminpu Group.


    Humerus: The preserved portion of the humerus (Figs. 2,3) is exposed in cranioventral view and measures 21.8 mm.The shaft is roughly circular in cross section and expandsmediolaterally to a width of about 5.9 mm across the epi-condyles a short distance proximal to the distal end. Proximalto the condyles, a shallow, triangular fossa embays the cranialsurface (Fig. 3) that tapers proximally. This fossa may behomologous with the brachial fossa, although it is shallowand smooth rather than a distinct scar like the brachial fossaof neornithean birds. The ellipsoidal dorsal and ventral con-dyles are subequal in size. The condyles lie immediately ad-jacent to one another, with only a shallow intercondylar sulcusbetween them. Both condyles are offset from the humerallong axis toward the ventral side of the humerus. Relative tothe longitudinal axis of the humerus, the ventral condyleparallels the entire humeral distal margin with its long axis

    Fig. 1. Locality (star) from which CAGS-IG-04-CM-023 was recovered near Changma, Gansu Province, Peoples Republic of China.

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    Harris et al. 549

    oriented distoventrallyproximodorsally. In contrast, the dorsalcondyle is oriented more horizontally, closer to perpendicularto the long axis of the humeral body. The dorsal condylealso sits in a slightly more proximal position than the ventral;its proximal margin thus invades the brachial fossa. Theventral condyle instead faces almost entirely distally. Theventral side of the distal end bears a large, circular, flat-topped but modestly rugose eminence in place of a typical,protuberance-style ventral epicondyle. Dorsal to the dorsalcondyle, the distal humerus is attenuated into a relativelylong process. At the dorsalmost visible margin, there is ashort, cranioventrally projecting ridge that demarcates the

    edge of a flat, craniodorsally facing, slightly elevated plat-form that may represent a dorsal epicondyle.

    Ulna: The 33.6 mm long ulna (Figs. 2, 3, 4A4C) appearsto have rotated slightly dorsally from its articulation with thehumerus and radius and is thus exposed in ventromedialview. Its shaft is bowed caudally for its proximal one-third,but straightens distal to that. The articular cotyle is triangularand canted proximocaudallydistocranially. The slightly con-cave articular surface is not obviously divided into distinctcotyles, though the accessory processes associated with suchcotyles still exist. The olecranon process consists