Cretornis Hlavaci Fric, 1881 from the Upper Cretaceous of Czech Republic

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Cretornis Hlavaci Fric, 1881 from the Upper Cretaceous of Czech Republic Cretaceous Research 55 (2015) 164e175 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes Cretornis hlavaci Fric, 1881 from the Upper Cretaceous of Czech Republic (Pterosauria, Azhdarchoidea) * Alexander Averianov a, b, , Boris Ekrt c a Zoological Institute of the Russian Academy of Sciences, Universitetskaya nab. 1, 199034 Saint Petersburg, Russia b Department of Sedimentary Geology, Geological Faculty, Saint Petersburg State University, 16 Liniya VO 29, 199178 Saint Petersburg, Russia c Department of Paleontology, National Museum Prague, Vaclavsk enam ĕstí 68, 115 79 Prague, Czech Republic article info abstract Article history: Cretornis hlavaci Fric, 1881 from the Upper Cretaceous (Turonian) of Czech Republic is a valid taxon Received 20 August 2014 referred to Azhdarchoidea based on having a saddle-shaped humeral head, pneumatic foramen on Accepted in revised form 27 February 2015 proximal humerus present on anterior side and absent on posterior side, elongate deltopectoral crest Available online 23 March 2015 with subparallel proximal and distal margins, pneumatic foramen absent on distal side of humerus, metacarpals IeIII not articulated with carpus and displaced on anterodorsal side of wing metacarpal, and Keywords: wing metacarpal much longer than humerus. Absence of a pneumatic foramen on posterior side of Pterosauria proximal humerus suggests attribution of Cretornis hlavaci to Neoazhdarchia. It has a unique construction Azhdarchoidea Cretornis of the distal ulna with a dorsal articulation surface placed distinctly proximal to the tuberculum shared Cretaceous only with the non-azhdarchid azhdarchoid Montanazhdarcho minor from the Campanian of North Czech Republic America. Cretornis hlavaci differs from the latter taxon by the structure of its humerus and distinctly longer wing metacarpal. It is more derived than “Tapejaridae” but shares with Azhdarchidae the del- topectoral crest of the humerus displaced distally from humeral head. Cretornis cannot be assigned to Azhdarchidae because of the oval cross section of the second wing phalanx. A unique rhombic outline of the distal humerus of Cretornis hlavaci is a possible autapomorphy for this taxon. Its wing span estimated as 1.5e1.6 m. This is the first taxon of non-azhdarchid pterosaurs known from the Upper Cretaceous of the Eastern Hemisphere. © 2015 Elsevier Ltd. All rights reserved. 1. Introduction short report attributed them to a new bird taxon named Cretornis Hlavaci (Fric, 1881a) but this lacked figures and a satisfactory Cretornis hlavaci Fric, 1881, one of the first Cretaceous pterosaurs description. The same year he published a short paper in Czech named from continental Europe and still the only doubtless accompanied with lithographic drawings (Fric, 1881b). It is not pterosaur record from the Czech Republic, was based on a partial possible to decide which paper has the priority. Here the Czech wing skeleton. The first fragments were found by Ms. Tomkovain paper (Fric, 1881b) is designated as the original description of 1880 and all subsequent fragments were collected by local phar- Cretornis hlavaci Fric, 1881 because it contains a more detailed macist Mr. Hlavac in a small quarry close to the village Zarecka description and illustrations. Fric identified the most complete Lhota near Chocen( ¼Chotzen in German papers) in the Kingdom of bone (humerus) as a coracoid and mentioned also the humerus Bohemia, part of the Austro-Hungarian Empire (now Czech Re- fragment and almost all forearm bones which are “astoundingly public). This discovery was studied by Antonín Fric (Anton Fritsch short proportionally to the coracoid” (Fric, 1881a:275). Later Fric in some German papers), a prominent Czech paleontologist (Ekrt, republished the figure in a book devoted to his studies of the fossils 2008). First he identified the bones as bird remains and, in a from the Bohemian Cretaceous (Fric, 1883: fig.45), but did not provide any additional description. This illustration is reproduced here (Fig. 1). In 1887 Fritsch presented the cast of Cretornis to the British Museum of Natural History in London and this specimen * Corresponding author. Zoological Institute of the Russian Academy of Sciences, was included in the “Catalogue of the Fossil Reptilia and Amphibia” Universitetskaya nab. 1, 199034 Saint Petersburg, Russia. of that museum (Lydekker, 1888). Lydekker (1988) was apparently E-mail addresses: [email protected] (A. Averianov), [email protected] fi (B. Ekrt). the rst to recognize the pterosaur nature of the specimen. He http://dx.doi.org/10.1016/j.cretres.2015.02.011 0195-6671/© 2015 Elsevier Ltd. All rights reserved. A. Averianov, B. Ekrt / Cretaceous Research 55 (2015) 164e175 165 Fig. 1. Cretornis hlavaci, holotype, left wing elements as illustrated by Fric (1883: fig.45). Zarecka Lhota, Czech Republic; Jizera Formation, Upper Cretaceous (Turonian). A, humerus in posterior view; B, humerus in anterior view; C, distal ulna in anterior view; D, proximal first wing phalanx in posterior view; E, proximal first wing phalanx in dorsal view; F, second wing phalanx in posterior view; G, distal wing metacarpal in anterior view. 166 A. Averianov, B. Ekrt / Cretaceous Research 55 (2015) 164e175 dismissed the name Cretornis and put the species within the genus these claims are incorrect. In azhdarchoids the ulnar crest is Ornithocheirus where all Cretaceous toothed pterosaurs were directed caudally (ventrally in flight position), similar to that in placed at that time. Lydekker changed the species epithet from ornithocheirids and pteranodontids. Bennett (1989: fig.2(6, 7)) hlavaci to hlavatschi following the German transliteration of this introduced this character based on comparison with the humerus Slavic name. However, there is no need for such emendation but the USNM 11925 (holotype of Bennettazhia oregonesis), where the ulnar Code requires diacritic marks to be deleted (ICZN, 1999:Article crest is not preserved. In the humerus of C. hlavaci deltopectoral 32.5.2.1), thus the correct species epithet would be hlavaci. crest is not “warped” and distal end is not triangular in distal view Lydekker (1988:14) noted that the humerus of the Czech specimen (see description and discussion below). “closely resembles the Wealden specimen” NHM UK 2353, the Institute abbreviations. IVPP e Institute of Vertebrate Paleon- holotype of Palaeornis cliftii Mantell, 1844, also referred by him to tology and Paleoanthropology, Beijing, China; NHM UK e National Ornithocheirus. NHM UK 2353 was recently reinterpreted as History Museum, London, United Kingdom; MOR e Museum of the belonging to a lonchodectid (Witton et al., 2009) or azhdarchoid Rockies, Bozeman, USA; NMP e National Museum Prague, Prague, (Averianov, 2012) pterosaur. Czech Republic; SMNK e Staatliches Museum für Naturkunde, Fric(Fritsch, 1905: fig.3) accepted Lydekker's interpretation of Karlsruhe, Germany; USNM e United States National Museum, the Czech fossil and reconstructed the wing of Ornithocheirus hla- Smithsonian Institution, Washington, USA. vaci and is reproduced here (Fig. 2). He thought that the bones Measurements of limb bones.Le length; PW e maximum width represented are humerus, proximal fragments of the ulna and of proximal end; DW e maximum width of distal end. All mea- radius, and first and second wing phalanges. The interpretation of surements are in millimeters. the humerus and wing phalanges is correct but the supposed proximal part of the radius is actually the distal part of the ulna and 2. Geologic setting proximal part of the ulna is the distal wing metacarpal (see description). New figures of the bones appeared in Fritsch and The holotype of Cretornis hlavaci Fric, 1881 was found at Zarecka Bayer (1905: figs. 32a, b, 33, pl.8, figs.1e8). Lhota, located in the eastern part of the Bohemian Cretaceous Basin Since the original reference by Lydekker (1988), Cretornis was (Fig. 3). This elongate basin extends from Saxony, through Bohemia either maintained as a synonym of Ornithocheirus (Kuhn, 1967), or and Moravia to Silesia. It represents shallow shelf sea with an ar- considered as Ornithocheiridae incertae sedis (Wellnhofer, 1978, chipelago of palaeo-highs. The most substantial were Central Eu- 1980; Barrett et al., 2008), or Pterosauria incertae sedis ropean Island (on the W, SW and S), West Sudetic Island (on the (Khozatsky and Yur'ev, 1964). More recently Averianov (2010) NW) and the East Sudetic Island (on the NE). Southeast part was noted that this forgotten taxon may be an azhdarchid. Here we adjacent with deep Tethys Ocean (Cech, 2011). Zarecka Lhota is provide detailed description and illustration of the fossils of Cre- approximately 30e40 km from the East Sudetic Island. Although tornis hlavaci and discuss the validity and taxonomic attribution of some smaller islands may have existed, there is no direct evidence this species of pterosaur. The orientation descriptors of the bones for this. A general lack of other terrestrial biota at Zarecka Lhota correspond to their position during flight with the wings suggests high sea stands, and as such, most of the palaeo-highs outstretched. would have been submerged. Jianu et al. (1997: non-paginated abstract) have studied the Cretornis was discovered in gray highly calcareous marlstones holotype of C. hlavaci and found that it is “clearly pteranodontid which were excavated in a small quarry ca 600 m west of the village based on having a caudally directed ulnar crest, a warped delto- Zarecka Lhota. Fric assigned the sequence to the Jizera beds (Iser- pectoral crest, and a triangular cross-section of the distal end.” All schichten in German, stratigraphic classification according Krejcí, Fig. 2. Cretornis hlavaci, reconstruction of the left wing of made by Fritsch (1905: fig.3) (A) and proposed here (B). In A individual bones have different orientation. In B the image is reversed and shows right wing in ventral view to make visible ulna which is ventral to radius in flight position. The proportions of wing elements in B are after Pteranodon (Bennett, 2001a: fig.1). 1, Humerus; 2, proximal radius in A and distal ulna in B; 3, proximal ulna in A and distal wing metacarpal in B; 4, first wing phalanx; 5, second wing phalanx.
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