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New Information on Postcranial Skeleton of the Early Cretaceous Gansus Yumenensis (Aves: Ornithuromorpha)

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New Information on Postcranial Skeleton of the Early Cretaceous Gansus Yumenensis (Aves: Ornithuromorpha) Historical Biology An International Journal of Paleobiology ISSN: 0891-2963 (Print) 1029-2381 (Online) Journal homepage: http://www.tandfonline.com/loi/ghbi20 New information on postcranial skeleton of the Early Cretaceous Gansus yumenensis (Aves: Ornithuromorpha) Ya-Ming Wang, Jingmai K. O'Connor, Da-Qing Li & Hai-Lu You To cite this article: Ya-Ming Wang, Jingmai K. O'Connor, Da-Qing Li & Hai-Lu You (2016) New information on postcranial skeleton of the Early Cretaceous Gansus yumenensis (Aves: Ornithuromorpha), Historical Biology, 28:5, 666-679, DOI: 10.1080/08912963.2015.1006217 To link to this article: http://dx.doi.org/10.1080/08912963.2015.1006217 View supplementary material Published online: 17 Feb 2015. Submit your article to this journal Article views: 130 View related articles View Crossmark data Citing articles: 2 View citing articles Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=ghbi20 Download by: [Shanghai Jiaotong University] Date: 08 October 2016, At: 19:48 Historical Biology, 2016 Vol. 28, No. 5, 666–679, http://dx.doi.org/10.1080/08912963.2015.1006217 New information on postcranial skeleton of the Early Cretaceous Gansus yumenensis (Aves: Ornithuromorpha) Ya-Ming Wanga,b*, Jingmai K. O’Connorb,c, Da-Qing Lid and Hai-Lu Youb,c aSchool of Earth Sciences and Resources, China University of Geosciences (Beijing), Beijing 100083, P.R. China; bKey Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Beijing 100044, P.R. China; cInstitute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, P.R. China; dGansu Geological Museum, Lanzhou, Gansu Province 730010, P.R. China (Received 13 December 2014; accepted 7 January 2015; first published online 17 February 2015) Gansus yumenensis Hou and Liu, 1984 is a basal ornithuromorph bird that has been recovered from the Lower Cretaceous Xiagou Formation in the Changma Basin of Gansu Province in northwestern China. Although it bears important significance in understanding the origin and early evolution of Ornithuromorpha, this taxon has only received preliminary morphological study. Here, we provide a detailed description of the postcranial morphology of Gansus yumenensis based on seven new specimens and new data from previously reported specimens, and update diagnosis for this taxon. Our phylogenetic analysis including new morphological data on Gansus yumenensis resolves all known four genera of Changma ornithuromorphs as more derived taxa than Jehol members of this clade, indicating the difference between Changma and Jehol avifauna. Keywords: Gansus yumenensis; Changma; Xiagou Formation; Early Cretaceous; Ornithuromorpha; anatomy; phylogeny Introduction 011, -07-CM-006, -05-CM-014, -04-CM-018 and -04-CM- Gansus yumenensis Hou and Liu, 1984 was named based 031), all from the Changma locality. We also describe some on an isolated left foot (IVPP V6862) collected from the details of two specimens (CAGS-IG-04-CM-003, -004) Lower Cretaceous Xiagou Formation in the Changma reported in You et al. (2006). They together provide a rare opportunity to study the detailed postcranial anatomy of Basin of Gansu Province in northwestern China. At the time of its discovery, this specimen represented the first basal ornithuromorphs. Notably, three of the new speci- Mesozoic fossil bird from China and the second oldest bird mens preserve gastroliths, contributing new information in the world, younger only than Archaeopteryx (Hou and about the habit of Gansus yumenensis. Liu 1984). Renewed field efforts in the Changma Basin nearly two decades later produced a wealth of far more complete specimens and You et al. (2006) reported five Institutional abbreviations specimens among them (CAGS-IG-04-CM-001 , 004, - CAGS-IG, Chinese Academy of Geological Sciences, 008), describing Gansus yumenensis as ‘a nearly modern Institute of Geology, Beijing; GSGM, Gansu Geological amphibious bird’. Their phylogenetic analysis resolved Museum, Lanzhou; IVPP, Institute of Vertebrate Paleontol- this taxon as a basal member of Ornithurae. Li et al. (2011) ogy and Paleoanthropology, Beijing. provided additional information on the morphology of the sternum, forelimb and hind limb based on nine fragmentary new specimens (IVPP V15074–15077, Systematic paleontology 15079–15081, 15083, 15084). In addition to Gansus, Aves Linnaeus, 1758 three other ornithuromorph taxa: Yumenornis huangi, Changmaornis houi and Jiuquanornis niui (Wang et al. Pygostylia Chiappe, 2002 2013), one enantiornithine Qiliania graffini (Ji et al. 2011) Ornithothoraces Chiappe and Calvo, 1994 and three unnamed enantiornithine specimens (You et al. Ornithuromorpha Chiappe, 2002 2005; Harris et al. 2006; Lamanna et al. 2006) have also Gansus yumenensis Hou and Liu, 1984 been described from the same locality. However, the vast majority of Changma avian specimens are referred to Gansus. Holotype specimen Here we describe seven new specimens of Gansus Distal left tibiotarsus and left tarsometatarsus with yumenensis (GSGM-07-CM-009, -07-CM-011, -06-CM- complete pedal digits (IVPP V6862; Hou and Liu 1984). *Corresponding author. Email: [email protected] q 2015 Taylor & Francis Historical Biology 667 Figure 1. Geographic location of the Changma Basin (indicated by star), Gansu Province, People’s Republic of China. Source: Modified from Wang et al. (2013). New referred specimens fenestrae; coracoid with a cranially hooked lateral process; GSGM-07-CM-009, articulated three caudalmost thoracic tibiotarsus long, with two strongly proximally projected vertebrae, synsacrum, pygostyle and partial pelvic girdle cnemial crests; position of metatarsal II trochlea high and consisting of right ilium, proximal pubis and ischium plantarly displaced relative to the metatarsal III trochlea; (Figure 2); GSGM-07-CM-011, articulated skeleton and pedal digit IV longest (autapormorphy); pedal claws with gastroliths, lacking the skull, cervical and caudal pointed flexor tubercles. vertebrae, pectoral girdles and forelimbs (Figure 3); GSGM-06-CM-011, nearly complete sternum, sternal ribs, partial furcula, sternal ribs and gastroliths (Figure 6); Description GSGM-07-CM-006, left proximal scapula, coracoid and Anatomical nomenclature primarily follows Baumel and partial left forelimb, with feather impression (Figure 7); Witmer (1993). English translations are used for GSGM-05-CM-014, complete left and partial right osteological terminology while Latin is maintained for forelimbs, complete hind limbs, scattered ribs and muscles. gastroliths (Figure 8); GSGM-04-CM-018, articulated left tibiotarsus and fibula (Figure 9); GSGM-04-CM-031, proximal and distal portions of left tarsometatarsus A (Figure 10). Locality and horizon In the vicinity of the Changma Township, Yumen City, Jiuquan area, Gansu Province, northwestern China; Lower Cretaceous (early Aptian), Xiagou Formation (You et al. 2005; Harris et al. 2006; Lamanna et al. 2006; Suarez et al. B 2013; Figure 1). ri syn il sp py ca dp Revised diagnosis is il tv pu lp Gansus yumenensis is distinguished from other known acf 1 cm ornithuromorphs by the following unique combination of characters: pygostyle narrow throughout length with Figure 2. (Colour online) Photograph (A) and line drawing (B) of Gansus yumenensis GSGM-07-CM-009. Abbreviations: acf, dorsal spinous ridge; sternum with well-developed acetabulum foramen; ca, caudal vertebrae; dp, dorsal process; il, craniolateral and lateral processes; lateral trabeculae of ilium; is, ischium; lp, lateral depression; pu, pubis; py, pygostyle; ri, sternum curved medially; sternum bearing a pair of caudal ridge; sp, spinous process; syn, synsacrum; tv, thoracic vertebrae. 668 Y.-M. Wang et al. Vertebral column and ribs (Figure 2), as in Apsaravis and Ichthyornis (Clarke and GSGM-07-CM-009 preserves the caudalmost three thor- Norell 2002; Clarke 2004); whereas the synsacrum is acic vertebrae in lateral view and GSGM-07-CM-011 formed by nine vertebrae in Yanornis and Yixianornis preserves two isolated thoracic vertebrae in caudal view. (Zhou and Zhang 2001; Clarke et al. 2006), nine or ten in The thoracic vertebrae possess well-developed rectangular Jianchangornis (Zhou et al. 2009), 11 in Changmaornis spinous processes, the dorsoventral height of which is (Wang et al. 2013) and only seven in the basal nearly half the total height of the associated vertebra Archaeorhynchus (possibly due to ontogenetic variation) (Figure 2). The transverse processes are long and robust (Zhou and Zhang 2006; Zhou et al. 2013). (CAGS-IG-04-CM-004; You et al. 2006). The vertebral The caudal series consists of seven free vertebrae, the foramen is large, nearly the same size as the articular cranial five of which possess robust transverse processes; surface of the centrum (Figure 3). The length of the centrum the processes become increasingly slender distally is markedly greater than the midpoint width (Figure 2). The (Figure 2). The pygostyle is short, slightly tapering to thoracic vertebrae appear to be amphicoelous. Like the end; its length is approximately equal to the combined enantiornithines, the lateral depressions on the centra are length of three free caudal vertebrae, as in CAGS-IG-04- developed as elongated grooves rather than broad fossae as CM-002 (You et al. 2006). The pygostyle is narrow in other ornithuromorphs (Figure 2). throughout its length and bears a spinous ridge along the Based on the number of transverse processes, the dorsal surface (Figure
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