Research 55 (2015) 164e175

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Cretaceous Research

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Cretornis hlavaci Fric, 1881 from the Upper Cretaceous of Czech Republic (Pterosauria, )

* 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 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 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 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 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 these claims are incorrect. In azhdarchoids the ulnar crest is 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, ; 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 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 (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. Scale bar equals 100 mm. A. Averianov, B. Ekrt / Cretaceous Research 55 (2015) 164e175 167

Diagnosis. A small pterodactyloid with estimated wing span 1.5e1.6 m referable to Azhdarchoidea based on the following combination of primitive () and derived (þ) characters: hu- meral head saddle-shaped (), pneumatic foramen on proximal face of the humerus present on anterior side (þ) and absent on posterior side (), elongate deltopectoral crest with subparallel proximal and distal margins (), pneumatic foramen absent on distal side of humerus (), metacarpals IeIII not articulated with carpus (þ) and displaced on anterodorsal side of wing meta- carpal (þ), and wing metacarpal much longer than humerus (þ). Differs from “Tapejaridae” and similar with Azhdarchidae by deltopectoral crest of humerus displaced distally from humeral head (þ). Further different from by the absence of pneumatic foramen on posterior side of proximal humerus (þ). Similar with non-azhdarchid azhdarchoid Mon- tanazhdarcho minor and different from all other Azhdarchoidea by dorsal articulation surface of distal ulna placed distinctly Fig. 3. A simplified geological map of the Bohemian Cretaceous Basin showing posi- proximal to tuberculum (þ), but otherwise different from that 0 0 tion of Zarecka Lhota locality (asterisk; N 49 59.59 ,E1614.26 ). A, Cretaceous rocks taxon by rhomboid shape of distal humerus (þ), lack of pneu- covered by younger Tertiary deposits; B, Cenomanian to middle Turonian deposits; C, matic foramen on distal humerus surface (), and distinctly upper Turonian to Coniacian deposits. Modified from Cech et al. (2005: fig.1A). longer wing metacarpal (þ). Differs from Azhdarchidae by oval cross section of second wing phalanx. Differs from other Pter- odactyloidea by rhomboid shape of distal humerus. fi 1869) and considered af liation to the Trigonia horizon (after Holotype. NMP Ob 10e0b 14, 0b 89eOb 92, Ob 101eOb 103, bivalve Trigonia) well developed in the west part of Bohemian incomplete left wing including humerus, distal fragment of ulna, Cretaceous Basin. Here, we refer the site to the Jizera Formation distal fragment of wing metacarpal, proximal fragment of first fi (regional lithostratigraphic classi cation after Cech et al., 1980). wing phalanx, and shaft fragment of second wing phalanx. However, the Jizera beds and Jizera Formation are diachronous Type locality and horizon.Zarecka Lhota, Czech Republic; Jizera fi (Cech et al., 1980) and it is very dif cult to correlate different parts Formation, Upper Cretaceous (middleeupper Turonian). of the basin due to the complex facial relationships. The Jizera Formation comprises mostly marlstones and silici- e clastic sandstones and is dated as middle late Turonian. It is 4. Description underlain by the loweremiddle Turonian BílaHoraFormation e and overlain by the upper Turonian lower Coniacian Teplice 4.1. Humerus Formation (Ulicný, 2001; Wiese et al., 2004). The Jizera Forma- e tion is a part of a transgressive regressive cycle deposited during The humerus is almost complete except the distal part of the the regressive phase after the maximum transgression charac- deltopectoral crest which is missing (Figs. 1A, B, 4). The bone sur- terized by the Bíla Hora Formation, which is composed of marl- face is abraded in some places. The humeral head is crescentic in stones, micritic limestones and spongolites. The vertebrate fauna proximal view, with shorter convex posterior margin and longer of the Jizera and Teplice formations, except the pterosaur Cre- concave anterior margin. The articular surface is saddle shaped, fi tornis, consist of diverse chondrichthyan and osteichthyan shes convex in anteroposterior section and concave in dorsoventral and marine reptiles (Fric, 1883; Ekrt et al., 2008; Kear et al., section. It is subdivided by a shallow groove into two unequal parts, 2014). smaller and more elevated dorsal part and a larger ventral part. This Detailed chronostratigraphy of the Zarecka Lhota site is unclear elevated part is quite distinct and is termed here the dorsal ridge of due to lack of index fossils and very distinct facies development of humeral head (Fig. 4A, B). The widest part of the articular surface in fi eastern part of basin compared to the signi cantly better explored proximal view is closer to the ventral side. The humeral neck is northwestern part (see Ulicný, 2001; Wiese et al., 2004). Based on inclined posteriorly to the long axis of the shaft at an angle of ~32. lithostratigraphy and position within the formation, most likely age The articular surface of the humeral head overhangs the neck along range if the site is upper part of the middle Turonian to the upper the posterior side. The anterior side of the neck and proximal shaft Turonian. is depressed between the ulnar and deltopectoral crests. Within Although the pterosaur bones are isolated, rock fragments with this depression, between the base of the deltopectoral crest and outer imprints indicate the wing skeleton was not in natural arti- humeral head, near the dorsal margin, there is a pneumatic fora- culated position, but the bones were disarticulated and accumu- men (Fig. 4A, C). lated more or less close together. Preserved surfaces of bones do not The deltopectoral crest is placed along the dorsal border of the show any traces of predator activity. The pterosaur remain is a humerus and directed anteriorly. It is separated from the proximal random occurrence accompanied with fortunate local rapid sedi- end by a relatively large distance: its proximal margin is located at mentation by silt and micrite. the level of the distal margin of the ulnar crest. The angle between the deltopectoral crest and the long axis of humeral head articu- 3. Systematic paleontology lation surface is ~133. The angle between the deltopectoral and ulnar crests in proximal view is ~88. Although the deltopectoral Pterosauria Kaup, 1834. crest is incomplete distally, its proportions suggest that it was not Azhdarchoidea Nesov, 1984. much longer than the ulnar crest. In the preserved part, the del- Cretornis Fric, 1881. topectoral crest is straight, without a sign of a distal curvature. Cretornis hlavaci Fric, 1881. The ulnar crest has a rounded apex and sloping proximal and Figs. 1, 2, 4e8 distal margins in anterior and posterior views. The apex is at the 168 A. Averianov, B. Ekrt / Cretaceous Research 55 (2015) 164e175 level distal to the most distal extension of the humeral head posterior of which is visible from the posterior side (Fig. 6B, C). articulation surface in posterior view. The apex of the crest is These grooves are apparently for the metacarpals IeIII and the somewhat deviated posteriorly. The articular surface of the hu- ventral bending of the metacarpal IV in this region is likely to meral head does not extend on the ulnar crest. provide a space for these reduced metacarpals. Location of the The narrowest point of the shaft is located immediately distal to attachment grooves of the metacarpals IeIII near the distal end of the distal margin of the deltopectoral crest. Its distal expansion is the metacarpal IV demonstrates that these bones were much more significant in anterior/posterior view compared to that in reduced in size compared with the wing metacarpal and certainly dorsal/ventral view. The shaft is round to oval in cross section. On did not articulate with the carpus. the posterior side of the shaft, proximal to the distal epiphysis, there is a prominent ridge-like supracondylar tubercle (Fig. 4C, E). 4.4. First wing phalanx The distal epiphysis is fully ossified and fused with the shaft. In distal view it is parallelogram-shaped with the roughly equal Approximately half of the shaft and the proximal end of this anteroventral/posterodorsal and anterodorsal/posteroventral sides bone is preserved (Figs. 1D, E, 7). There is also a more proximal (Fig. 4D). The pointed ectepicondyle and entepicondyle form fragment of the shaft which contact with the larger fragment is respectively the dorsal and ventral ends of this parallelogram. On missing. On the original Fric drawings, the pieces are connected the anterior side of the distal epiphysis there is a prominent together, but they are not adjacent and therefore were unconnected bulbous capitulum (radial condyle, articulating also with the by us. The shaft is oval in cross-section and convex anteriorly in capitular cotyle of ulna). The capitulum is separated from the crest- dorsal and ventral views. In anterior and posterior views the shaft is like ectepicondyle by a deep groove. A similarly deep groove (fovea slightly depressed ventrally in the place of anterior convexity. The supratrochlearis) is placed along the proximal margin of the proximal end is somewhat abraded and the posterior process of the capitulum. The long axis of the capitulum is inclined at the angle of proximal end is missing. The extensor tendon process is also ~48 to the long axis of the shaft in anterior view. The trochlea incomplete but undoubtedly it was fully fused with the bone. The (ulnar condyle articulating with the trochlear cotyle of ulna) is of cotyles articulating with the pulley-shaped distal condyle of the peculiar morphology. It is represented by a ridge parallel to the wing metacarpal are separated by a strong ridge. The dorsal cotyle distal surface of the distal epiphysis and a round depression adja- is about twice narrower compared with the ventral condyle and cent distally to this ridge. A similar but much smaller depression on certainly would be about twice as long if the posterior process the bulbous trochlea is present in adult specimens of Pteranodon would be preserved. The intercotylar groove is shallow and has a (Bennett, 2001a: fig.74) and Azhdarcho lancicollis (Averianov, 2010: slight depression where should be a pneumatic foramen. Because of fig.24). The central part of the distal surface of the distal epiphysis is poor preservation of the bone surface it is unclear if the pneumatic slightly concave but there is no a pneumatic foramen there. There is foramen was really present there. a poorly developed and low ulnar tubercle posterior to this depression (Fig. 4D). 4.5. Second wing phalanx

4.2. Ulna The second wing phalanx is represented by most of the shaft but proximal and distal ends are missing (Figs. 1F, 8). The shaft is oval in The fragment preserves approximately more than half of the cross-section and slightly expanded towards the ends. It is convex shaft and the distal end (Figs. 1C, 5). The shaft is oval in cross- anteriorly as the first wing phalanx. Compared to Fric assemblage, section and has the same width in dorsal and ventral views. In we were able to insert several other related fragments and now the anterior and posterior views it is only little expanded towards the bone is rather longer than on the original Fric drawings. distal end. The distal end is quite peculiar in morphology. It is dominated by the tuberculum forming the larger ventral side of the 4.6. Measurements distal end. The tuberculum has a well-defined and rounded artic- ular surface. The most unusual feature of this bone is that the Humerus: L ¼ 75.6; PW ¼ 16.7; DW ¼ 25.6. Ulna: DW ¼ 14.9. tuberculum is projecting far more distally compared with the smaller dorsal articular surface. On the anterior side of the bone 4.7. Wing span estimation there are two parallel lines probably representing the muscle scars (Figs. 1C, 5B) possibly for m. flexor digitorum longus (Bennett, The estimation of the wing span in life for pterosaurs is a diffi- 2008: fig.2A). The ventral most of these lines is longer and ex- cult task because it should take into consideration the natural tends on to the tuberculum. A similar muscle scar can be found on flexures of the elbow, carpus, and wing finger (Bennett, 2001b). For the ulna of Pteranodon (Bennett, 2001a: fig.75A) and Azhdarchidae wing-span estimation we used several published skeletal re- (Godfrey and Currie, 2005: fig.16.7D). This scar is possibly for the constructions of pterodactyloids in flight (Table 1). These mea- ventral ulno-carpal ligament (Bennett, 2008: fig.2A). surements give the following linear regression (1) and exponential fit (2) between the estimated wing span (WS) and humerus length 4.3. Metacarpals (HL) calculated by the scatterplot module of STATISTICA 7.1©Stat- Soft, Inc.: The wing metacarpal is represented by approximately half of the shaft and the distal end which is mostly concealed within the piece WS ¼ 35:32 þ 20:46 HL (1) of rock (Figs. 1G, 6). The shaft is oval in cross-section with flattened or slightly depressed anterior side. In anterior and posterior views WS ¼ 874:38 expð0:0072 HLÞ (2) the part of shaft before the distal end is slightly deviated ventrally but the distal end is parallel to the main part of the shaft. In dorsal According to these equations, the estimated wing span of Cre- and ventral views the shaft is slightly sinusoid with the anterior tornis hlavaci with humerus length of 75.6 mm would be curvature near the distal end and the posterior curvature some 1582.01 mm (1) or 1506.94 mm (2), which is roughly 1.51e1.58 m. distance proximal to the latter. On the dorsal side, near the ventral Because the holotype is a fully mature specimen, this wing span bend of the shaft, there are three short parallel grooves, the most estimate is likely the maximum for the species. A. Averianov, B. Ekrt / Cretaceous Research 55 (2015) 164e175 169 170 A. Averianov, B. Ekrt / Cretaceous Research 55 (2015) 164e175

Fig. 6. Cretornis hlavaci, holotype, distal fragment of left wing metacarpal in ventral Fig. 5. Cretornis hlavaci, holotype, distal fragment of left ulna in ventral (A), anterior (A), posterior (B), dorsal (C), and anterior (D) views. The distal condyle is concealed in (B), dorsal (C), posterior (D), and distal (E) views. Zarecka Lhota, Czech Republic; Jizera the rock matrix. Zarecka Lhota, Czech Republic; Jizera Formation, Upper Cretaceous Formation, Upper Cretaceous (Turonian). Abbreviations: das, dorsal articulation sur- (Turonian). Abbreviations: dcn, dorsal condyle; mcIeIII, grooves for attachment of face; tub, tuberculum. Scale bar equals 10 mm. metacarpals IeIII; vcn, ventral condyle. Scale bar equals 10 mm.

5. Phylogenetic position of Cretornis hlavaci Bakonydraco galaczi; this clade is the sister to Azhdarchidae. The last result is intriguing because of the geographic and stratigraphic The material of Cretornis hlavaci was coded for 11 from 111 proximity of Cretornis hlavaci and Bakonydraco galaczi but it is characters (9.9%) from the pterosaur data matrix provided by certainly an artifact of tree calculating because these taxa have no Andres and Ji (2008). These characters are 84(1), 85(0), 86(1), 87(5), overlapping codings. The same analysis of the matrix after 88(1), 89(1), 90(0), 91(0), 93(0), 100(2), and 106(0). The new removing of the unstable Bakonydraco galaczi produces two trees technology search algorithm of TNT (Goloboff et al., 2003) with with the length of 326 steps, consistency index of 0.49, and default settings produced three most parsimonious trees with the retention index of 0.81. Cretornis hlavaci appears on both trees as a length of 327 steps, consistency index of 0.48, and retention index sister taxon for Azhdarchidae. This result should be considered of 0.80. Cretornis hlavaci appears on these trees as 1) a sister taxon with some caution, however, because the data matrix analyzed for Azhdarchidae including Bakonydraco galaczi from the Santonian includes limited number of azhdarchoid taxa and distribution of } of Hungary (Osi et al., 2005, 2011); 2) a sister taxon for Azhdarch- some characters is poorly understood in taxa known from flattened idae while Bakonydraco galaczi is the sister taxon for the clade skeletons. On the other hand, in taxa known from fragmentary but Chaoyangopteridae þ Azhdarchidae; 3) a sister taxon for three dimensional materials, like Cretornis hlavaci, the proportions

Fig. 4. Cretornis hlavaci, holotype, left humerus in proximal (A), posterior (B), anterior (C), distal (D), dorsal (E), and ventral (F) views. All photos are stereopairs. Zarecka Lhota, Czech Republic; Jizera Formation, Upper Cretaceous (Turonian). Abbreviations: cap, capitulum; dc, deltopectoral crest; dr, dorsal ridge of humeral head; ect, ectepicondyle; ent, ente- picondyle; h, humeral head; pf, pneumatic foramen; st, supracondylar tubercle; tro, trochlea; uc, ulnar crest; ut, ulnar tubercle. Scale bar equals 10 mm. A. Averianov, B. Ekrt / Cretaceous Research 55 (2015) 164e175 171

Fig. 8. Cretornis hlavaci, holotype, shaft fragment of left second wing phalanx in dor- sal(?) (A), posterior (B), and ventral(?) (C) views. Zarecka Lhota, Czech Republic; Jizera Formation, Upper Cretaceous (Turonian). Scale bar equals 10 mm.

Table 1 Humerus length and wing span in life calculated from published skeletal re- constructions of pterodactyloids in flight.

Taxon Humerus Wing Reference length, mm span, mm

Aurorazhdarcho 62.4 1024 Frey et al., 2011: fig.6a primordius gracilis 108 2723 Brower, 1983: fig.1A fi Fig. 7. Cretornis hlavaci, holotype, proximal fragment of left rst wing phalanx in santanae 208 4176 Wellnhofer, 1991: fig.24 proximal (A), dorsal (B), posterior (C), ventral (D), and anterior (E) views. Zarecka Santanadactylus spp 240 4700 Wellnhofer, 1985: fig.48a Lhota, Czech Republic; Jizera Formation, Upper Cretaceous (Turonian). A small piece of Anhanguera piscator 255 4998 Kellner and Tomida, 2000: fig.3a a more distal shaft fragment has no contact with the main fragment. Abbreviations: dc, Pteranodon spp 269 5959 Bennett, 2001b: fig.124a dorsal cotyle; vc, ventral cotyle. Scale bar equals 10 mm. a Scale is not provided for the reconstruction. Wing span is calculated from the humerus length given in the text. 172 A. Averianov, B. Ekrt / Cretaceous Research 55 (2015) 164e175 of the wing elements and other limb bones cannot be calculated. 5.4. Position of the deltopectoral crest on humeral shaft Here we provide discussion of some phylogenetically important characters of Cretornis hlavaci which may further facilitate its In most pterodactyloids the deltopectoral crest of humerus is taxonomic attribution. positioned proximally, close or confluent to the humeral head. The crest is considerably displaced distally from the proximal end in fi 5.1. Humeral head shape and dorsal ridge Nyctosauridae (Williston, 1903: pl.42, g.7) and to a somewhat less extent in Azhdarchidae (Godfrey and Currie, 2005: fig.16.6C, D; fi The humeral head is convex in both anteroposterior and Averianov, 2010: g.23) and non-azhdarchid azhdarchoids (Witton fi dorsoventral sections in Istiodactylidae (Hooley, 1913: pl.39, fig.2) et al., 2009: gs 3D, E, 4D, E, 6D, E, 7D, E) including USNM 11925, and Ornithocheiridae (Wellnhofer, 1991: fig.17; Hazlehurst and the holotype of Bennettazhia oregonensis from the Albian of Oregon, Rayner, 1992; Kellner and Tomida, 2000: figs 30, 31; Veldmeijer, USA (personal observation by AA). In Dsungaripteridae (IVPP “ ” fi 2003: figs 15, 16). It is saddle-shaped, convex in anteroposterior V4059, V4062) and Tapejaridae (Eck et al., 2011: g.7, pl.1; Aires fi section and concave in dorsoventral section in et al., 2014: g.6) the deltopectoral crest has the primitive prox- (Bennett, 2001a: fig.70), Nyctosauridae (Williston, 1903: pl.42, imal position. The distal displacement of deltopectoral crest in fig.7), Dsungaripteridae (IVPP V4059, V4062), “Tapejaridae” (Eck Nyctosauridae and non-tapejarid azhdarchoids was likely derived fi et al., 2011: fig.7, pl.1; Aires et al., 2014: fig.6), non-azhdarchid in parallel. The deltopectoral crest in Cretornis hlavaci ts this azhdarchoids (Witton et al., 2009: figs 3, 4, 6, 7), and Azhdarchi- derived condition of non-tapejarid Azhdarchoidea. dae (Averianov, 2010: fig.23). This difference in the shape of hu- meral head apparently reflects specialization of the flight 5.5. Deltopectoral crest of humerus shape mechanism in Ornithocheiroidea sensu stricto (excluding Pter- anodontidae and Nyctosauridae, which evidently do not belong to In pterodactyloids the deltopectoral crest of the humerus is quite that group; Hazlehurst and Rayner, 1992). The humeral head in diverse in shape. In pteranodontids the deltopectoral crest has a Cretornis hlavaci is saddle-shaped which is consistent with its long base and subparallel (converging) proximal and distal margins, attribution to Azhdarchoidea. The dorsal ridge of humeral head is with the distal end is rolled ventrally and oriented obliquely to the fi prominent, but not hypertrophied as in Thalassodrominae (Aires humerus long axis (Bennett, 2001a: gs 69, 70). In Ornithocheiridae et al., 2014: fig.6). and Istiodactylidae the distal margin of the deltopectoral crest is perpendicular to the humerus long axis while the proximal margin slopes towards the distal end of the crest which is oriented 5.2. Pneumatic foramen on anterior side of proximal humerus perpendicular to the long axis of the humerus (Hooley, 1913: pl.39, fig.2; Wellnhofer,1991: fig.17; Kellner and Tomida, 2000: figs 30, 31; The pneumatic foramen on the anterior side of proximal hu- Veldmeijer, 2003: fig.15). These constructions are quite different merus between the deltopectoral crest and humeral head is present and so-called “warped” deltopectoral crest (Unwin, 2003) should in Cretornis hlavaci (Fig. 4A, C), Azhdarchidae (Godfrey and Currie, not be considered as a synapomorphy for Ornithocheiroidea. The fi fi 2005: g.16.6D, E; Averianov, 2010: g.23D, I; Buffetaut et al., construction of the deltopectoral crest in Pteranodon is more similar fi 2011: g.1D), non-azhdarchid azhdarchoids (Witton et al., 2009: to the primitive pterodactyloid condition and in the earliest pter- fi fi “ ” gs 3D, 4D, 6A, 7A; Andres and Myers, 2013: g.3), Tapejaridae anodontian Alamodactylus byrdi from the Coniacian of North fi fi (Eck et al., 2011: g.7, pl.1B; Aires et al., 2014: g. 6B, E), Pter- America it is almost unwarped (Myers, 2010a: fig.2; Andres and fi anodontidae (Bennett, 2001a: g.70B), and Nyctosauridae (Bennett, Myers, 2013: fig.4A, B), suggested that the “warped” condition in 1989: 673). This foramen is notably absent in Dsungaripteridae, the Pteranodon and ornithocheirids was developed in parallel. Under likely sister group for Azhdarchoidea (Unwin, 2003; Lü et al., 2010). this character optimization the hatchet-shaped deltopectoral crest Thus, presence of this pneumatic foramen could be a synapomor- of Nyctosaurus (Williston, 1903: pl.42, fig.7) is not a reversal from phy for either Azhdarchoidea, under character optimization the “warped” condition but is a modification of a primitive pter- preferred parallelisms, or more inclusive clade, if reversals are odactyloid construction. The proximal humerus fragment (USNM preferred. 13804) from the Turonian strata of Eagle Ford Group in Texas, USA, was referred initially to Pteranodon by Gilmore (1935) and to Pter- 5.3. Pneumatic foramen on posterior side of proximal humerus anodontidae by Bennett (1989: fig.2(1e4)), who treated the family in a broad sense, including Ornithocheirus. This specimen has the A large pneumatic foramen is present on the posterior side of ornithocheiroid construction of the deltopectoral crest and may the humerus near the proximal end and on the level between the belong to the Ornithocheiridae, which are known from the Cen- humeral head and ulnar crest in Ornithocheiridae (Bennett, 1989: omanian strata of the Eagle Ford Group (Myers, 2010b; Andres and figs 1(4), 2(1); Wellnhofer, 1991: fig.17e; Kellner and Tomida, Myers, 2013). The humerus with mostly missing deltopectoral crest 2000: fig.31b, c; Veldmeijer, 2003: fig.15D). This foramen is ab- from the Lower Cretaceous of Peru (Bennett, 1989: fig.1) also likely sent in Pteranodontidae and Nyctosauridae (Bennett, 1989, belongs to Ornithocheiridae. In Dsungaripteridae (IVPP V4059, 2001a; Unwin, 2003), as well as in non-azhdarchid azhdarch- V4062), “Tapejaridae” (Eck et al., 2011: fig.7, pl.1; Aires et al., 2014: oids (Witton et al., 2009: figs 3, 4, 6, 7), including Mon- fig.6), and Azhdarchidae (Godfrey and Currie, 2005: fig.16.6CeE; tanazhdarcho minor (MOR 691), and in Azhdarchidae (Godfrey Averianov, 2010: fig.23) the deltopectoral crest is elongate, with and Currie, 2005: fig.16.6D; Averianov, 2010: fig.23). In Dsun- subparallel proximal and distal margins, which is likely the primi- garipteridae this character is variable, the foramen may be pre- tive morphology for pterodactyloids. In Cretornis hlavaci the delto- sent (IVPP V4059) or absent (IVPP V4062). In “Tapejaridae,” pectoral crest has this construction. which is likely paraphyletic, the pneumatic foramen on posterior side of proximal humerus is present in Tapejara (Eck et al., 2011: 5.6. Shape of distal end of humerus fig.7: pl.1D) but absent in (Aires et al., 2014: fig.6A, D). This foramen is absent in Cretornis hlavaci which suggest its In Istiodactylidae and Ornithocheiridae the distal end of hu- placement in the clade Neoazhdarchia (Tupuxuara þ merus is more or less in the form of an isosceles triangle, with the Azhdarchidae). anterior side longest (Hooley, 1913: pl.39, fig.3; Wellnhofer, 1985: A. Averianov, B. Ekrt / Cretaceous Research 55 (2015) 164e175 173

fig.8f; Kellner and Tomida, 2000: fig.32c, d; Veldmeijer, 2003: uniting these taxa, or has a broader distribution among Azh- fig.15F; Averianov et al., 2005: fig.2E). The exact shape of the distal darchoidea. The humerus morphology is different in these two end of humerus is not known for Pteranodon because all available taxa. In any case, the distal ulna morphology is consistent with specimens are crushed, but it is likely subtriangular (Bennett, placing Cretornis hlavaci within Azhdarchoidea. 2001a: 75). This derived condition is usually considered as the synapomorphy of Ornithocheiroidea (Unwin, 2003). The distal end 5.9. Metacarpals IeIII articulation with carpus of humerus is oval to D-shaped, with somewhat flattened anterior side, in Nyctosaurus (coding in Andres and Ji, 2008), Dsungar- In basal pterodactyloids the metacarpals IeIII are long and ipteridae (IVPP V4059, V4062), “Tapejaridae” (Rodrigues et al., articulate with the carpus (Wellnhofer, 1970: fig.4E, F; Andres and 2011: fig.8B), non-azhdarchid azhdarchoids (Witton et al., 2009: Ji, 2008: fig.4A). In sinensis according to the original figs 6B, 7B), and Azhdarchidae (Godfrey and Currie, 2005: fig.16.6G; description “only metacarpal I and IV articulate with the carpus” Averianov, 2010: fig. 24C; Rodrigues et al., 2011: fig. 8A, C, D). The (Andres and Ji, 2006: 75, fig.4C, D). However, metacarpals were shape of distal end of humerus in Cretornis hlavaci (Fig. 4D) fits obviously confused in that paper and the longest metacarpal, nearly neither of these conditions. It is of rhomboid shape, or more reaching the carpus, is the metacarpal III. Subsequently Andres and correctly, of parallelogram shape, because the sides are not equal. Ji (2008) coded this taxon as having metacarpal III articulating with This morphology has not been described for any other pter- the carpus and metacarpals I and II reduced. In Ornithocheiridae odactyloid and it is considered here as an autapomorphy of Cre- metacarpal III is quite stout and long and reaches the carpal region, tornis hlavaci. while metacarpals I and II are short and do not articulate with the carpus (Wellnhofer, 1991: figs 18, 31a, b, 38a, b). Metacarpals IeIII 5.7. Pneumatic foramen on distal end of humerus are greatly reduced in length, do not articulate with the carpus, and are placed at the distal end of metacarpal IV in Pteranodontidae A large and round pneumatic foramen is present on distal end of (Bennett, 2001a: fig.88) and “Tapejaridae” (Wang and Zhou, 2003: humerus in Istiodactylidae (Hooley, 1913: pl.39, fig.3) and Orni- fig.1). In Nyctosaurus the digits I to III may be totally lost (Bennett, thocheiridae (Wellnhofer, 1985: fig.8f; Kellner and Tomida, 2000: 2000), although a small metacarpal was previously described for fig.32c, d; Veldmeijer, 2003: fig.15F; Averianov et al., 2005: fig.2E). that taxon (Williston, 1903: 145, pl.42, fig.6). Andres and Ji (2008) In Dsungaripteridae a small pneumatic foramen could be present coded Dsungaripteridae as having metacarpals IeIII articulated (IVPP V4059) or absent (IVPP V4062). In “Tapejaridae,” non- with the carpus but according to Lü et al. (2009: 639) “metacarpals azhdarchid azhdarchoids, and Azhdarchidae such foramen is ab- IeIII do not reach the distal carpals, but they occupy approximately sent, but some much smaller pneumatic foramina or fenestration half the length of the metacarpal IV.” In IVPP V4062 the longest of might be present (Godfrey and Currie, 2005: fig.16.6G; Witton et al., these metacarpals, apparently metacarpal III, is nearly reaching the 2009: figs 6B, 7B; Averianov, 2010: fig.24C; Rodrigues et al., 2011: proximal end of metacarpal IV but not the carpus, although this fig.8). A small ventral pneumatic foramen on distal humerus sur- bone may be incomplete proximally in this specimen. Possibly in face is present in non-azhdarchid azhdarchoid Radiodactylus lang- Dsungaripteridae condition of this character was similar to that in stoni from the Albian of Texas, USA (Murry et al., 1991; Andres and Istiodactylidae, with long metacarpal III articulating or nearly Myers, 2013). A distinct cleft-like pneumatic foramen is present on reaching the carpus, and reduced metacarpals I and II. Azhdarchi- the distal end of the humerus in Montanazhdarcho minor dae were coded by derived state of this characters, with short (McGowen et al., 2002 and personal observation of MOR 691 by AA) distally placed metacarpals IeIII not articulating with the carpus from the Campanian of Montana, USA, originally described as (Andres and Ji, 2008), which is likely but currently not supported by azhdarchid but reinterpreted as non-azhdarchid azhdarchoid description of the material. In Tapejara the metacarpals IeIII are (Averianov, 2014). In Cretornis hlavaci there is no pneumatic fora- incomplete proximally, but the metacarpal III is much more robust men on the distal end of humerus, which is consistent with the compared with metacarpals I and II (Eck et al., 2011: fig.8, pl.2), azhdarchoid condition. which may suggest that it was also longer. According to Unwin (2003: 170) in Tapejara at least one of metacarpals IeIII retains 5.8. Morphology of distal ulna end contact with the carpus but he did not specify the specimen upon which this observation is based. In SMNK PAL 380, the postcranial In most pterodactyloids the distal end of ulna has quite uniform skeleton from Crato Formation of Brazil referable to “Tapejaridae” morphology, with the dorsal articulation surface at the same level (Unwin and Martill, 2007: fig.17.11a, c), one of the medial meta- or only little proximal to the ventral fovea and a distinct tuberc- carpals, likely metacarpal III, is distinctly longer than two others ulum (Hooley, 1913: pl.39, fig.6; Bennett, 2001a: fig.75A; Godfrey metacarpals (I and II) and may nearly reach the carpus. In Sinop- and Currie, 2005: fig.16.7C, D; Averianov, 2010: fig.25FeJ). The terus dongi the metacarpals IeIII are short and clearly do not reach distal ulna morphology in Cretornis hlavaci is distinct from that the carpus (Wang and Zhou, 2003). In Cretornis hlavaci the meta- construction. Here the ventral fovea is not differentiated from the carpals IeIII were reduced in length and confined to the distal end tuberculum and the dorsal articulation surface is placed far more of the metacarpal IV as inferred from their attachment grooves proximally compared with the proximal end of tuberculum. The (Fig. 6B, C). Obviously these metacarpals do not articulate with the only other pterodactyloid which has such unusual construction of carpus. the distal ulna is Montanazhdarcho minor (MOR 691), a non- azhdarchid azhdarchoid (Averianov, 2014). The tuberculum and 5.10. Position of metacarpals IeIII relative to wing finger ventral fovea projects distally at some extent relative to the dorsal metacarpal articulation surfaces in Ornithocheiridae (Wellnhofer, 1991: fig.18a) and “Tapejaridae” (Eck et al., 2011: fig.7, pl.2; the tuberculum is Initially in pterodactyloids metacarpals IeIII lay in their inadequately termed “olecranon” in that paper, the olecranon is the anatomical position, medial to the metacarpal IV, i.e. on its anterior process on the proximal end of the ulna). The morphology of distal side in flight position (Wellnhofer, 1970: fig. 4E, F). This primitive ulna end is poorly known in azhdarchoids and it is unclear if the condition preserved in Istiodactylidae (Andres and Ji, 2006: fig.1), unusual structure of this region in Cretornis hlavaci and Mon- Ornithocheiridae (Wellnhofer, 1991: figs 31a, b, 38a, b), and tanazhdarcho minor represents a synapomorphy for the group Dsungaripteridae (IVPP V4062). In Pteranodontidae metacarpals 174 A. Averianov, B. Ekrt / Cretaceous Research 55 (2015) 164e175

IeIII are displaced on the anterodorsal side of the distal end of (Averianov, 2014). A similar but much less pronounced ridge could metacarpal IV (Bennett, 2001a: figs 88, 93), which is likely corre- be present in Dsungaripteridae (IVPP V4062). The second wing lated with changing in the flexing plane of the manual digits IeIII phalanx of Cretornis hlavaci (Fig. 8) is oval in cross-section and lack (Bennett, 2001a: 90). Cretornis hlavaci has this derived condition, such a ridge, which safely rule out attribution of this taxon to with metacarpals IeIII on the anterodorsal side of metacarpal IV Azhdarchidae. (Fig. 6B, C). The condition of this character in “Tapejaridae” and Azhdarchidae is unclear. 6. Conclusions

5.11. Wing metacarpal length The cladogram in Fig. 9 summarizes the phylogenetic position of Cretornis hlavaci based on distribution of characters which could be The wing metacarpal, short in non-pterodactyloid pterosaurs, is considered as unambiguous synapomorphies and which are known at least 0.8 the humerus length in pterodactyloids, which is in all the included terminal taxa. There are two more characters considered as one of the synapomorphies for the group (Unwin, which are potentially important but their distribution is not known “ ” e 2003). According to the coding in Andres and Ji (2008) the wing for Tapejaridae : 1) position of wing metacarpals I III relative to metacarpal is between 0.67 and 1.81 humerus length in Istio- wing metacarpal and 2) wing metacarpal length. The wing meta- dactylidae, Ornithocheiridae, Chaoyangopteridae, and “Tapejar- carpal is relatively short in and Tapejara (see above) and e idae,” and more than 1.81 in Pteranodontidae, Nyctosauridae, these taxa also likely have metacarpals I III in anatomical position, Dsungaripteridae, and Azhdarchidae. In Montanazhdarcho minor anterior to the wing metacarpal. If condition of these characters is this ratio is only 1.47 (McGowen et al., 2002: tab.1), similar to this more derived in Tupuxuara, this will further elaborate the concept ratio (1.6) in Sinopterus dongi (Wang and Zhou, 2003: tab.2). In of Neoazhdarchia (Unwin, 2003). Cretornis hlavaci now can be fi Cretornis hlavaci the preserved fragment of wing metacarpal is con dently considered as non-azhdarchid azhdarchoid and a fi 95 mm in length, which is greater than the humerus length member of the clade Neoazhdarchia (Fig. 9). This is the rst taxon of (75.6 mm), but the preserved portion has no sign of proximal non-azhdarchid azhdarchoids known from the Upper Cretaceous of expansion suggesting that almost half of the bone is missing. The the Eastern Hemisphere. complete bone was certainly longer than 1.81 of the humerus length. The gracile preserved distal part of wing metacarpal is Acknowledgments markedly different from the stout wing metacarpal in Mon- tanazhdarcho minor (McGowen et al., 2002: fig.1) and “Tapejaridae” AA thanks J. Shunxing (Institute of Vertebrate Paleontology and (Eck et al., 2011: fig.8, pl.3). Paleoanthropology, Beijing, China), J. Scanella (Museum of the Rockies, Bozeman, USA), Eberhard Frey (Staatliches Museum für Naturkunde, Karlsruhe, Germany), and H.-D. Sues (United States 5.12. Second and third wing phalanges cross section National Museum, Smithsonian Institution, Washington, USA) for access to the pterosaur specimens. We are grateful to D. Grigoriev In Azhdarchidae the second and third phalanges of wing digit (Saint Petersburg State University, Saint Petersburg, Russia) for are distinctly T-shaped in cross section, with a longitudinal ridge taking photographs. We thank Eberhard Frey (Staatliches Museum along its ventral surface (Martill and Frey, 1999: fig.2; Averianov, für Naturkunde, Karlsruhe, Germany), David Martill (School of 2007: pl.8, fig.2; Averianov, 2010: fig.32A, B). A supposed tape- Earth and Environmental Sciences, University of Portsmouth, Great jarid specimen with T-shaped wing phalanges from the Lower Britain), and an anonymous reviewer for reviewing the paper. The Cretaceous Crato Formation of Brazil belongs to Azhdarchidae laboratory work of AA was supported by the Russian Scientific Fund project 14-14-00015. BE thanks the Ministry of Culture of Czech Republic (MKCR) for support (project nr. DF12P01OVV021) and Z. Staffen for advice.

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