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Laser-stimulated fluorescence (LSF) has seen increased use in palaeontological investigations in recent years. The method uses the high flux of laser light of visible wavelengths to reveal details sometimes missed by traditional long-wave ultraviolet (UV) methods using a lamp. In this study, we compare the results of LSF with UV-A-generated fluorescence on a range of fossils from the Upper Jurassic Solnhofen Limestone Konservat-Lagerstätte of Bavaria, Germany. The methodology follows previous protocols of LSF with modifications made to enhance laser beam intensity, namely keeping the laser at a constant distance from the specimen, using a camera track. Our experiments show that along with making surface details more vivid than UV-A or revealing them for the first time, LSF has the additional value of revealing shallow subsurface specimen detail. Fossil decapods from the Solnhofen Limestone reveal full body outlines, even under the matrix, along with details of segmentation within the appendages such as limbs and antennae. The results indicate that LSF can be used on invertebrate fossils along with vertebrates and may often surpass the information provided by traditional UV methods.

Brighstoneus gen. nov. Etymology. Brighstoneus is named after the village of Brighstone on the Isle of Wight, which is close to the excavation site and was home to the Reverend William Fox, a celebrated Victorian fossil collector whose discoveries had a major impact on early dinosaurian research. Type species. Brighstoneus simmondsi gen. et sp. nov. Diagnosis. As for type and only species (see below). Locality and horizon. Wessex Formation, early Barremian, Lower Cretaceous. MIWG 6344 was excavated during 1978, from a plant debris bed (L9 of Stewart 1978) to the west of Grange Chine on the south coast of the Isle of Wight (Fig. 2). Comment on association. The skeleton was found associated with one of the UK��� s most complete theropods, Neovenator salerii Hutt et al., 1996 (MIWG 6348; Brusatte et al. 2008). Both individuals were contained, with some overlap, in an area of ~ 3 �� 6 m. There was no articulated material, but preservation was consistent across individual elements and there was no replication of material or other iguanodontian material found during the excavation. The material was stored at the Museum of the Isle of Wight Geology, who were also involved in the excavation. The then curator is confident that the material was associated (S. Hutt, pers. comm. 2021). Some photographs and drawings of the site are included in Supplementary material (Figs S1���S5) and hard copies of other contemporaneous records have been accessioned under MIWG 6344. Comment on stratigraphy. The base of the Barremian stage within the Wessex Formation lies west of Sudmoor Point and has been dated to 126.5 Ma (Gale et al. 2020), while the Barremian���Aptian boundary at the base of Chron M0, on the basis of dates from Svalbard (Zhang et al. 2019), is given as 121.4 Ma (Gale et al. 2020). This gives the Barremian a duration of approximately 5.1 Ma compared to earlier estimates of 4.5 Ma based on phosphorus burial rates (Bodin et al. 2006). The Mantellisaurus atherfieldensis holotype was found following a cliff fall from the Shepherds Chine Member of the Vectis Formation in 1914 (Hooley 1925) and was probably from earliest Aptian strata, making it approximately 4.0 Ma younger than Brighstoneus simmondsi, assuming uniform depositional rates., Published as part of Lockwood, Jeremy A. F., Martill, David M. & Maidment, Susannah C. R., 2021, A new hadrosauriform dinosaur from the Wessex Formation, Wealden Group (Early Cretaceous), of the Isle of Wight, southern England, pp. 847-888 in Journal of Systematic Palaeontology 19 (12) on pages 3-4, DOI: 10.1080/14772019.2021.1978005, http://zenodo.org/record/5696995, {"references":["Stewart, D. J. 1978. The sedimentology and palaeoenvironment of the Wealden Group of the Isle of Wight, southern England. Unpublished PhD thesis, University of Portsmouth, 346 pp.","Hutt, S., Martill, D. M. & Barker, M. J. 1996. The first European allosauroid dinosaur (Lower Cretaceous, Wealden Group, England). Neues Jahrbuch fur Geologie und Palaontologie, Monatshefte, 10, 635 - 644.","Brusatte, S. L., Benson, R. B. J. & Hutt, S. 2008. The osteology of Neovenator salerii (Dinosauria: Theropoda) from the Wealden Group (Barremian) of the Isle of Wight. Palaeontographical Society Monographs, 162, 1 - 75 t 45 pls.","Allen, P. & Wimbledon, W. A. 1991. Correlation of NW European Purbeck-Wealden (non-marine Lower Cretaceous) as seen from the English type-areas. Cretaceous Research, 12, 511 - 526.","Sweetman, S. C. 2007. Aspects of the microvertebrate fauna of the Early Cretaceous (Barremian) Wessex Formation of the Isle of Wight, southern England. Unpublished PhD thesis, University of Portsmouth, 316 pp. https: // pure. port. ac. uk / ws / portalfiles / portal / 6060512 / SCS _ 2007 _ PhD _ Thesis. pdf","Gale, A. S., Mutterlose, J., Batenburg, S., Gradstein, F. M., Agterberg, F. P., Ogg, J. G. & Petrizzo, M. R. 2020. The Cretaceous Period. Pp. 1 - 63 in F. M. Gradstein, J. G. Ogg, M. D. Schmitz & G. M. Ogg (eds) Geologic time scale 2020. Volume 2. Elsevier, Amsterdam.","Zhang, Y., Ogg, J. G., Minguez, D. A., Hounslow, M., Olaussen, S., Gradstein, F. M. & Esmeray Senlet, S. 2019. Magnetostratigraphy of U / Pb-dated boreholes in Svalbard, Norway, implies that the Barremian - Aptian boundary (beginning of Chron M 0 r) is 121.260. 4 Ma. AGU Annual Meeting (San Francisco, 9 - 13 December 2019). GP 44 A- 06. https: // agu. confex. com / agu / fm 19 / meetingapp. cgi / Paper / 577991.","Bodin, S., Godet, A., F ¨ ollmi, K. B., Vermeulen, J., Arnaud, H. & Strasser, A. 2006. The late Hauterivian Faraoni oceanic anoxic event in the western Tethys: evidence from phosphorus burial rates. Palaeogeography, Palaeoclimatology, Palaeoecology, 235, 245 - 264.","Hooley, R. W. 1925. On the skeleton of Iguanodon atherfieldensis sp. nov., from the Wealden Shales of Atherfield (Isle of Wight). Quarterly Journal of the Geological Society of London, 81, 1 - 61."]}

Lockwood, Jeremy A. F., Martill, David M., Maidment, Susannah C. R. (2021): A new hadrosauriform dinosaur from the Wessex Formation, Wealden Group (Early Cretaceous), of the Isle of Wight, southern England. Journal of Systematic Palaeontology 19 (12): 847-888, DOI: 10.1080/14772019.2021.1978005

Brighstoneus simmondsi gen. et sp. nov . (Figs 3���22, 24) Etymology. The specific name honours Mr Keith Simmonds who made the discovery of the specimen. Holotype. MIWG 6344, a partial skeleton composed of the following elements: dorsal process of right premaxilla; both maxillae; both jugals; left palpebral; left nasal; both dentaries; predentary; one transitional dorsal and seven dorsal vertebrae; sacrum; six caudal vertebrae; dorsal ribs, nine from the left side and five from the right side; both ilia; right ischium; possible prepubic process; and the right femur. Some parts of the same individual (including two dorsal vertebrae and other fragments) remain in private ownership and are not described herein. Diagnosis. Differs from all other iguanodontians by possessing the following autapomorphies and unique combination of characters (autapomorphies indicated with an asterisk): maxillary crowns possessing both a primary ridge and mesially placed accessory ridges on the lingual surface *; nasal expanded postnarially to produce a modest nasal bulla with convex lateral walls *. Acharacter combination of at least 28 dentary tooth positions in a dentary with one active crown and one replacement tooth for each position and non-parallel alveolar septa. In addition, Brighstoneus can be distinguished from other Barremian���Aptian Wealden Group iguanodontians by possession of the following combination of features: ratio of precoronoid length of the dentary to minimum depth> 6.0; coronoid process projects at approximately 90 �� with respect to the dorsal margin of dentary; bilobed ���heart-shaped��� ventral predentary process with prominent anterior denticles with concave mesial and distal edges; posteriorly positioned maxillary ascending process in lateral view with length of the anterior section approximately twice the length of the posterior section; prominent anterodorsal process present on maxilla; anterior (maxillary) process of jugal relatively long (60% of overall length) and tapers distally to form a triangular ending; ventral section of the posterior margin of the jugal (heel) projects posteriorly to form a spurlike feature; in dorsal view the jugal is straight; ventral border overlapped by maxillary process of premaxilla; anteroventral nasal process forms posteroventral margin of external narial opening; ventral surface of ischiadic peduncle of ilium parallel with the anteroventral margin of the postacetabular process; ischiadic peduncle of ilium has flat lateral wall with no pronounced posterolateral boss; ventral shelf at base of iliac preacetabular process weakly developed; dorsoventrally deep preacetabular process with little or no axial twist; deep and short iliac central plate with ratio of depth to length> 1.2. Description Overall, MIWG 6344 is excellently preserved with little distortion or crushing (measurements are available in the Supplementary material). Many of the bones exhibit unusual eroded areas, which have left smooth, ���scooped out��� regions extending down to and including the internal trabecular bone. Some of these areas are partially covered by the original sediment, suggesting these features formed prior to, or soon after, burial. It is unlikely that this damage resulted from prolonged subaerial exposure as the cortical surface is generally well preserved (Behrensmeyer 1978). As many of these eroded features are associated with highly cartilaginous regions, it is possible that they are examples of invertebrate bioerosion, potentially by isopterans (Britt et al. 2008, and references therein; Huchet et al. 2011), which are known from the Wealden Group (Jarzembowski 1981). The term ���eroded��� is used in the descriptive text to indicate these areas. There is also evidence, particularly on the ribs, of shallow circular pits varying between 2���4 mm in diameter, which might represent dermestid beetle activity (Britt et al. 2008). Premaxilla Only the dorsal (nasal) process of the right premaxilla is preserved, but without its posterior-most tip (Fig. 3). In lateral view, the fragment has a long mediolaterally thin process, which is gently curved, convex dorsally and gradually tapers as it extends posteriorly. The lateral surface is rounded and grooved in the posterior threefifths of its length for contact with the anterodorsal process of the nasal (Fig. 3A: this area is described as ���bevelled��� in Mantellisaurus atherfieldensis rather than grooved: Norman 1986). Anteriorly the ventral margin curves ventrally forming a thin blade of bone, which is incomplete but would have contributed to the internarial septum. The medial surface of the dorsal process is flat for articulation with its antimere. Maxilla Both maxillae are present, with the left being the better preserved. The left maxilla (Fig. 4) is almost complete, although it has lost parts of the anterior ends of the anteroventral and anterodorsal processes. Between the anterior processes a fragment of bone has been glued into a position that is probably anatomically incorrect (Fig. 4A, C). The ascending process is almost complete, having a thin fractured margin, but preserving the groove for reception of the maxillary process of the premaxilla. The maxilla has been subjected to minor transverse lateral compression that has slightly relocated the incomplete jugal process medially. The maxilla is a robust bone but is relatively thinner in dorsal and ventral views than that of Mantellisaurus atherfieldensis (NHMUK PV R5764). Its general shape in lateral view is of an anteroposteriorly expanded triangle. The apex is directed dorsally as the ascending process but is situated in a markedly posterior position, and if complete the section anterior to this process would have been approximately twice the length of the posterior section. Aratio of ~ 2.0 or higher is also seen in Altirhinus kurzanovi (2.1: Norman 1998), Zhanghenglong yangchengensis (2.3: Xing et al. 2014), Protohadros byrdi (2.3: Head 1998), Proa valdearinnoensis (2.4: McDonald et al. 2012a) and Ouranosaurus nigeriensis (2.6: Taquet 1976). In contrast the ratio is 1.4 in Mantellisaurus atherfieldensis (NHMUK PV R5764). In lateral view the ventral edge of the maxilla is shallowly concave, the tooth row being almost straight for most of its length. There are 29 tooth positions: most of the alveoli are empty but two established and two emerging replacement crowns are present (see below). In ventral view the tooth row is straight for most of its length but posteriorly the last quarter curves laterally. The bone preservation of the lateral surface is better in the right maxilla, which has eight small nutrient foramina (1���3 mm in diameter) that open anterolaterally and form an anteroposteriorly orientated row in the ventral half of the maxilla. Anteriorly, the ventral surface of the maxilla forms an anteroventral process that, although incomplete, follows the curve of the tooth row and tapers anteriorly in lateral view. The first alveolus is situated at the base of this process, anterior to which the process is edentulous. Although the area dorsal to the anteroventral process has suffered some damage, a prominent anterodorsal process is present. Posteriorly the transverse cross-section of this process is triangular with a gently dorsoventrally concave lateral surface, a flat ventromedial surface and a dorsomedial surface that is roughened and has a longitudinal groove in its posterior section. Extending anteriorly, the process becomes increasingly compressed transversely to form a thin blade, whose end is missing. Possession of an anterodorsal process is considered ancestral for Archosauria (Wagner & Lehman 2009) and has been employed as a character in iguanodontian phylogeny (e.g. Sues & Averianov 2009; Xing et al. 2014; McDonald et al. 2017). Its presence has been used to distinguish Saurolophinae from Lambeosaurinae (Wagner & Lehman 2009), but it is also present in non-hadrosaurids such as Altirhinus kurzanovi (Norman 1998), Bactrosaurus johnsoni (Godefroit et al. 1998), Eolambia caroljonesa (CEUM 35492: McDonald et al. 2012b) and Gilmoreosaurus mongoliensis (Prieto-Marquez & Norell 2010), and is especially prominent in Protohadros byrdi (Head 1998). The anterior and posterior dorsal edges of the lateral wall of the maxilla meet to form the apex of the transversely compressed ascending process. The jugal process arises from the base of the posterior edge of the ascending process (Fig. 4A, D). The trough between the jugal process and lateral body of the maxilla expands posteriorly to form a mediolaterally broad ectopterygoid shelf. When viewed dorsally the posterior maxilla is more ovoid (long axis anteroposterior) and the medial wall more convex than in Mantellisaurus atherfieldensis (NHMUK PV R5764). Below the apex of the ascending process and situated posteriorly is a depressed area, which would presumably have contributed to the antorbital fenestra and might have formed a small antorbital fossa (Fig. 4A). This depressed area is also present in Iguanodon bernissartensis (Norman 1980), Mantellisaurus atherfieldensis (NHMUK PV R5764), Ouranosaurus nigeriensis (Taquet 1976), Camptosaurus dispar (Gilmore 1909) and Bolong yixianensis (Wu & Godefroit 2012). There is no evidence of an antorbital fenestra in Altirhinus kurzanovi (Norman 1998) or Jinzhousaurus yangi (Wang & Xu 2001; Barrett et al. 2009) and this feature is not described in Equijubus normani (McDonald et al. 2014). Anteriorly, the lateral part of the dorsal surface is furrowed to form the premaxillary groove for contact with the premaxilla (Fig. 4A, B). Medially the maxilla has a vertical surface that has an elongate rectangular outline with the ascending process rising above its dorsal edge. Above the tooth row is a concave arcade of circular special foramina (sensu Edmund 1957) or replacement foramina (sensu Jin et al. 2010), the curve of which is considerably deeper than the ventral edge (Fig. 4C). Between the special foramina and the tooth row the bone is thin with a textured surface and forms the alveolar parapet. Dorsally there is a medial shelf that is damaged anteriorly but appears to have a rounded edge. Nasal Only the left nasal is preserved (Fig. 5). It is undistorted but broken at both ends, so missing the posterior articulation for the frontal and prefrontal. The nasal is elongate anteroposteriorly, relatively broad and convexly curved mediolaterally so that with its antimere the external surface formed a vault over the nasal cavity. Anteriorly the nasal is divided into two processes. The anterodorsal process is dorsoventrally deeper in lateral view than the anteroventral process. The anterodorsal section, which articulated with the dorsal process of the premaxilla, is incomplete but the area that contributed to the posterior margin of the external naris is intact. The base of the anteroventral process is present showing that the nasal also made a contribution to the posteroventral margin of the external naris. There is no anteroventral process in Iguanodon bernissartensis (Norman 1980), Mantellisaurus atherfieldensis (Norman 1986), Protohadros byrdi (Head 1998), Bactrosaurus johnsoni (Godefroit et al. 1998), Equijubus normani (McDonald et al. 2014) or Gobihadros mongoliensis (Tsogtbaatar et al. 2019). In hadrosaurids an anteroventral process is usually present, for example in Maiasaura peeblesorum and Gryposaurus notabilis (Prieto-Marquez & Norell 2010) and is particularly prominent in Brachylophosaurus canadensis (Prieto-Marquez 2001) and Edmontosaurus annectens (Campione & Evans 2011). Among non-hadrosaurid hadrosauriforms, an abbreviated anteroventral process is present in Altirhinus kurzanovi (Norman 1998), with longer processes in Ouranosaurus nigeriensis (Taquet 1976), Jinzhousaurus yangi (Wang & Xu 2001; Barrett et al. 2009) and Bolong yixianensis (Wu & Godefroit 2012). The posterior margin of the external naris is almost straight and posteroventrally orientated. The ventral border of the lateral surface of the nasal has a shallowly bevelled surface (Fig. 5C, E) and presumably underlapped the maxillary process of the premaxilla as a scarf joint, although anteriorly there is also a slight groove along the dorsal edge of this surface. The nasals of Gobihadros mongoliensis and Choyrodon barsboldi have a grooved ventrolateral surface (Tsogtbaatar et al. 2019) and both appear to underlap the posterolateral process of the premaxilla; Bactrosaurus johnsoni (Godefroit et al. 1998) and Altirhinus kurzanovi (Norman 1998) are grooved along their ventral borders. The medial border of the dorsal surface is grooved anteriorly in Brighstoneus simmondsi, presumably where it articulated with the dorsal process of the premaxilla, but tapers to a thin blade posteriorly that most likely formed a simple butt joint with its antimere (see Weishampel 1984) as in Altirhinus kurzanovi (Norman 1998) and Jinzhousaurus yangi (Barrett et al. 2009), although in Ouranosaurus nigeriensis the nasals are asymmetrical with one having a blade and the other a groove along the dorsal edge (Taquet 1976). When viewed laterally in life position the dorsal border of the nasal is sinusoidal and forms a distinct anterior convexity with the most dorsal part of the curve situated posterior to the posterior border of the external naris. The section of the lateral wall of the nasal, posterior to the naris, is slightly convex in ventral view. Collectively these features would have produced a rounded bulge and formed a distinct step between the anterior and posterior sections of the nasal (Fig. 5E). This is less prominent but similar in shape and position to the nasal bulla of the iguanodontian Muttaburrasaurus langdoni (Bartholomai & Molnar 1981), although in Muttaburrasaurus the lateral walls are slightly concave. Ahigh arched region, dorsal to the posterior narial, is also seen in Altirhinus kurzanovi (Norman 1998), Choyrodon barsboldi (Gates et al. 2018) and in some hadrosaurids, for example Gryposaurus notabilis (Lambe 1914) and Kritosaurus navajovius (Brown 1910). Ouranosaurus nigeriensis also has a rounded protuberance, but this is situated more posteriorly on both nasals, just anterior to the suture for the frontals (Taquet 1976), while Jinzhousaurus yangi has mediolaterally convex nasals with a sagittal depression between them (Barrett et al. 2009), a feature also seen in Altirhinus kurzanovi (Norman 1998) and Choyrodon barsboldi (Gates et al. 2018). There is no evidence of a sagittal trough in Brighstoneus simmondsi. Temporal and geographical separation, together with the pattern of phylogenetic relationships and variety of morphologies in these examples, suggests that nasal ornamentation evolved independently on several occasions in iguanodontids. This feature also appears to be unique among the Wealden Group iguanodontids. Jugal Both jugals are preserved and are almost complete (Fig. 6). The dorsal sections of the postorbital process and the process overlapping the quadratojugal are broken. It is a triradiate bone with an anteriorly directed maxillary process, a central dorsally projecting postorbital process and a posterior dorsally projecting quadratojugal process. Both jugals are very straight and narrow in dorsal view (Fig. 6E, F) compared to Mantellisaurus atherfieldensis (NHMUK PV R5764), which is laterally convex with a relatively wider orbital shelf. The maxillary process in lateral view occupies 60% of the total length of the jugal (ratio of length anterior to midpoint of postorbital process/maximum length �� 0.60). This ratio varies considerably in iguanodontians, from 0.29 in Xuwulong yueluni (You et al. 2011) and 0.33 in Iguanodon bernissartensis (Norman 1980) to 0.56 in Mantellisaurus atherfieldensis (NHMUK PV R 5764) and 0.62 in Ouranosaurus nigeriensis (Taquet 1976). However, in most hadrosauriforms including hadrosaurids this ratio ranges between 0.40 and 0.54. For example, Altirhinus kurszanovi (0.46: Norman 1998), RBINS R57 (0.52: Norman 1986), Equijubus normani (0.49: You et al. 2003), Eolambia caroljonesa (0.48: McDonald et al. 2012b), Brachylophosaurus canadensis (0.51: Prieto-Marquez 2001) and Gryposaurus latidens (0.48: Prieto-Marquez 2012). The maxillary process in lateral view is gently curved with a convex ventral margin and a concave dorsal margin. The process is quite gracile and lacks the dorsoventral expansion seen in many hadrosauriforms, such as those present in Probactrosaurus gobiensis (Norman 2002); Eolambia caroljonesa (McDonald et al. 2012b) and Gobihadros mongoliensis (Tsogtbaatar et al. 2019), instead the anterior dorsal margin slopes anteroventrally to join the ventral margin as a point. This slope forms a facet for articulation with the lacrimal. Amaxillary process with a pointed end, lacking dorsoventral expansion, is also seen in Altirhinus kurzanovi (Norman 1998), Batyrosaurus rozhdestvenskyi (Godefroit et al. 2012), Dakotadon lakotaensis (Weishampel & Bjork 1989) and Choyrodon barsboldi (Gates et al. 2018). The maxillary process of the jugal in Mantellisaurus atherfieldensis (NHMUK PV R 5764) is not dorsoventrally expanded but has a much blunter, less tapered anterior end, albeit with some damage ventrally. Medially the maxillary process is deeply excavated to form an anteroposteriorly elongate facet with a roughened surface for reception of the finger-like jugal process of the maxilla. The jugal is expanded medially at the posterior end of the maxillary facet to form a buttress (Fig. 6E, F) for the distal end of the jugal process of the maxilla. Posterior to the maxillary facet is a posterodorsally orientated ridge that separates the facet from a shallow oval fossa for the ectopterygoid. Between the maxillary and postorbital processes, the dorsal surface is concave in lateral view and provided a shelf, forming the ventral margin of the orbit. The orbit is bounded posteriorly by the postorbital process, which is long, straight and ascends perpendicular to the main axis of the jugal body. Dorsally it develops an anterolaterally facing facet for suture with a reciprocal facet on the jugal process of the postorbital. Between the postorbital process and the quadratojugal process is a ���U���-shaped embayment that forms the ventral border of the lateral temporal fenestra. On its medial surface, the posterior and ventral margins of the fenestra have a subtle, smooth and shallowly convex rolled edge (Fig. 6B, C), which is not readily apparent in other hadrosauriforms, including Mantellisaurus atherfieldensis (NHMUK PV R 5764), Eolambia caroljonesa (McDonald et al. 2012b) and Sirindhorna khoratensis (Shibata et al. 2015). The right jugal has a nutrient foramen ~ 1 mm in diameter, located medially below the ventral border of the fenestra and postorbital process, while on the left jugal two foramina are present (with d, Published as part of Lockwood, Jeremy A. F., Martill, David M. & Maidment, Susannah C. R., 2021, A new hadrosauriform dinosaur from the Wessex Formation, Wealden Group (Early Cretaceous), of the Isle of Wight, southern England, pp. 847-888 in Journal of Systematic Palaeontology 19 (12) on pages 5-31, DOI: 10.1080/14772019.2021.1978005, http://zenodo.org/record/5696995, {"references":["Behrensmeyer, A. K. 1978. Taphonomic and ecologic information from bone weathering. Paleobiology, 4, 150 - 162.","Britt, B. B., Scheetz, R. D. & Dangerfield, A. 2008. A suite of dermestid beetle traces on dinosaur bone from the Upper Jurassic Morrison Formation, Wyoming, USA. Ichnos, 15, 59 - 71.","Huchet, J. B., Deverley, D., Gutierrez, B. & Chauchat, C. 2011. Taphonomic evidence of a human skeleton gnawed by termites in a Moche-civilisation grave at Huaca de la Luna, Peru. International Journal of Osteoarchaeology, 21, 92 - 102.","Jarzembowski, E. A. 1981. An Early Cretaceous termite from southern England (Isoptera: Hodotermitidae). Systematic Entomology, 6, 91 - 96.","Norman, D. B. 1986. On the anatomy of Iguanodon atherfieldensis (Ornithischia: Ornithopoda). Bulletin de L'institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre, 56, 281 - 372.","Norman, D. B. 1998. On Asian ornithopods (Dinosauria, Ornithischia). 3. A new species of iguanodontid dinosaur. Zoological Journal of the Linnean Society, 122, 291 - 348.","Xing, H., Wang, D., Han, F., Sullivan, C., Ma, Q., He, Y., Hone, D. W. E., Yan, R., Du, F. & Xu, X. 2014. A new basal hadrosauroid dinosaur (Dinosauria: Ornithopoda) with transitional features from the Late Cretaceous of Henan Province, China. PLoS ONE, 9, e 98821. doi: 10. 1371 / journal. pone. 0098821","Head, J. J. 1998. A new species of basal hadrosaurid (Dinosauria, Ornithopoda) from the Cenomanian of Texas. Journal of Vertebrate Paleontology, 18, 718 - 738.","McDonald, A. T., Espilez, E., Mampel, L, Kirkland, J. I. & Alcala, L. 2012 a. An unusual new basal iguanodont (Dinosauria: Ornithopoda) from the Lower Cretaceous of Teruel, Spain. Zootaxa, 3595, 61 - 76.","Taquet, P. 1976. Geologie et paleontologie du gisement de Gadoufaoua (Aptien du Niger). Cahiers de paleontologie. Editions du Centre National de la Recherche Scientifique, Paris, 191 pp.","Wagner, J. R. & Lehman, T. M. 2009. An enigmatic new lambeosaurine hadrosaur (Reptilia: Dinosauria) from the Upper Shale member of the Campanian Aguja Formation of Trans-Pecos Texas. Journal of Vertebrate Paleontology, 29, 605 - 611.","Sues, H. - D. & Averianov, A. 2009. A new basal hadrosauroid dinosaur from the Late Cretaceous of Uzbekistan and the early radiation of duck-billed dinosaurs. Proceedings of the Royal Society B, 276, 2549 - 2555.","McDonald, A. T., Gates, T. A., Zanno, L. E. & Makovicky, P. J. 2017. Anatomy, taphonomy, and phylogenetic implications of a new specimen of Eolambia caroljonesa (Dinosauria: Ornithopoda) from the Cedar Mountain Formation, Utah, USA. PLoS ONE, 12, 176896. doi: 10. 1371 / journal. pone. 0176896","Godefroit, P., Dong, Z. - M., Bultynck, P., Li, H. & Feng, L. 1998. Sino-Belgian Cooperative Program. Cretaceous dinosaurs and mammals from Inner Mongolia: 1) New Bactrosaurus (Dinosauria: Hadrosauroidea) material from Iren Dabasu (Inner Mongolia, P. R. China). Bulletin de l' Institute Royal des Sciences Naturelles du Belgique, 68, 1 - 70.","McDonald, A. T., Bird, J., Kirkland, J. I. & Dodson, P. 2012 b. Osteology of the basal hadrosauroid Eolambia caroljonesa (Dinosauria: Ornithopoda) from the Cedar Mountain Formation of Utah. PLoS ONE, 7, e 45712. doi: 10.1371 / journal. pone. 0045712","Prieto-Marquez, A. & Norell, M. A. 2010. Anatomy and relationships of Gilmoreosaurus mongoliensis (Dinosauria: Hadrosauroidea) from the Late Cretaceous of Central Asia. American Museum Novitiates, 3694, 1 - 52.","Norman, D. B. 1980. On the ornithischian dinosaur Iguanodon bernissartensis of Bernissart (Belgium). Memoires de l'Institut Royal des Sciences Naturelles de Belgique, 178, 1 - 105.","Gilmore, C. W. 1909. Osteology of the Jurassic reptile Camptosaurus, with a revision of the species of the genus, and descriptions of two new species. Proceedings of the United States National Museum, 36, 197 - 332.","Wu, W. & Godefroit, P. 2012. Anatomy and relationships of Bolong yixianensis, an Early Cretaceous iguanodontoid dinosaur from Western Liaoning, China. Pp. 292 - 333 in P. Godefroit (ed.) Bernissart dinosaurs and Early Cretaceous terrestrial ecosystems, Indiana University Press, Bloomington and Indianapolis.","Wang, X. & Xu, X. 2001. A new iguanodontid (Jinzhousaurus yangi gen. et sp. nov.) from the Yixian Formation of western Liaoning, China. Chinese Science Bulletin, 46, 1669 - 1672.","Barrett, P. M., Butler, R. J., Wang, X. - L. & Xu, X. 2009. Cranial anatomy of the iguanodontoid ornithopod Jinzhousaurus yangi from the Lower Cretaceous Yixian Formation of China. Acta Palaeontologica Polonica, 54, 35 - 48.","McDonald, A. T., Maidment, S. C. R., Barrett, P. M., You, H. - L. & Dodson, P. 2014. Osteology of the basal Hadrosauroid Equijubus normani (Dinosauria, Ornithopoda) from the Early Cretaceous of China. Pp. 44 - 72 in D. A. Eberth & D. C. Evans (eds) Hadrosaurs. Indiana University Press, Bloomington and Indianapolis.","Edmund, A. G. 1957. On the special foramina in the jaws of many ornithischian dinosaurs. Contributions of the Royal Ontario Museum, 48, 3 - 14.","Jin, L. - Y., Chen, J., Zan, S. - Q., Butler, R. J. & Godefroit, P. 2010. Cranial anatomy of the small ornithischian dinosaur Changchunsaurus parvus from the Quantou Formation (Cretaceous: Aptian - Cenomanian) of Jilin Province, northeastern China. Journal of Vertebrate Paleontology, 30, 196 - 214.","Tsogtbaatar, K., Weishampel, D. B., Evans, D. C. & Watabe, M. 2019. A new hadrosauroid (Dinosauria: Ornithopoda) from the Late Cretaceous Baynshire Formation of the Gobi Desert (Mongolia). PLoS ONE, 14, e 0208480. doi: 10.1371 / journal. pone. 0208480","Campione, N. E. & Evans, D. C. 2011. Cranial growth and variation in Edmontosaurus (Dinosauria: Hadrosauridae): implications for latest Cretaceous megaherbivore diversity in North America. PLoS ONE, 6, e 25186. doi: 10.1371 / journal. pone. 0025186","Weishampel, D. B. 1984. Evolution of jaw mechanisms in ornithopod dinosaurs. Advances in Anatomy, Embryology and Cell Biology, 87, 1 - 59.","Bartholomai, A. & Molnar, R. E. 1981. Muttaburrasaurus, a new iguanodontid (Ornithischia: Ornithopoda) dinosaur from the Lower Cretaceous of Queensland. Memoirs of the Queensland Museum, 20, 319 - 49.","Gates, T. A., Tsogtbaatar, K., Zanno, L. E., Chinzorig, T. & Watabe, M. 2018. A new iguanodontian (Dinosauria: Ornithopoda) from the Early Cretaceous of Mongolia. PeerJ, 6, e 5300. doi: 10.7717 / peerj. 5300","Lambe, L. M. 1914. On Gryposaurus notabilis, a new genus and species of trachodont dinosaur from the Belly River Formation of Alberta, with a description of the skull of Chasmosaurus belli. The Ottawa Naturalist, 27, 145 - 155.","Brown, B. 1910. The Cretaceous Ojo Alamo Beds of New Mexico with description of the new dinosaur genus Kritosaurus. Bulletin of the American Museum of Natural History, 28, 267 - 274.","You, H., Li, D. & Lui, W. 2011. A new hadrosauriform Dinosaur from the Early Cretaceous of Gansu Province, China. Acta Geologica Sinica, 85, 51 - 57.","You, H., Ji, Q., Li, J. & Li. Y. 2003. A new hadrosauroid dinosaur from the mid-Cretaceous of Liaoning, China. Acta Geologica Sinica, 77, 148 - 154.","Prieto-Marquez, A. 2001. Osteology and variation of Brachylophosaurus canadensis (Dinosauria, Hadrosauridae) from the Upper Cretaceous Judith River Formation of Montana. Unpublished Master' s thesis, Montana State University, Bozeman, 390 pp.","Prieto-Marquez, A. 2012. The skull and appendicular skeleton of Gryposaurus latidens, a saurolophine hadrosaurid (Dinosauria: Ornithopoda) from the early Campanian (Cretaceous) of Montana, USA. Canadian Journal of Earth Sciences, 49, 510 - 532.","Norman, D. B. 2002. On Asian ornithopods (Dinosauria: Ornithischia). 4. Probactrosaurus Rozhdestvensky, 1966. Zoological Journal of the Linnean Society, 136, 113 - 144.","Godefroit, P., Escuille, F., Bolotsky, Y. L. & Lauters, P. 2012. A new basal hadrosauroid dinosaur from the Upper Cretaceous of Kazakhstan. Pp. 335 - 362 in P. Godefroit (ed.) Bernissart dinosaurs and Early Cretaceous terrestrial ecosystems. Indiana University Press, Bloomington and Indianapolis.","Weishampel, D. B. & Bjork, P. R. 1989. The first indisputable remains of Iguanodon (Ornithischia: Ornithopoda) from North America: Iguanodon lakotaensis, sp. nov. Journal of Vertebrate Paleontology, 9, 56 - 66.","Shibata, M. & Azuma, Y. 2015. New basal hadrosauroid (Dinosauria: Ornithopoda) from the Lower Cretaceous Kitadani Formation, Fukui, central Japan. Zootaxa, 3914, 421 - 440.","Weishampel, D. B., Jianu, C. - M., Csiki, Z. & Norman, D. B. 2003. Osteology and phylogeny of Zalmoxes (n. g.), an unusual euornithopod dinosaur from the latest Cretaceous of Romania. Journal of Systematic Palaeontology, 1, 65 - 123.","Godefroit, P., Codrea, V. & Weishampel, D. B. 2009. Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda), based on new specimens from the Upper Cretaceous of Nalat-Vad (Romania). Geodiversitas, 31, 525 - 553.","Dalla Vecchia, F. M. 2009. Tethyshadros insularis, a new hadrosauroid dinosaur (Ornithischia) from the Upper Cretaceous of Italy. Journal of Vertebrate Paleontology, 29, 1100 - 1116.","Maryanska, T. & Osmolska, H. 1979. Aspects of hadrosaurian cranial anatomy. Lethaia, 12, 265 - 273.","Prieto-Marquez, A., Erickson, G. M. & Ebersole, J. A. 2016. Anatomy and osteohistology of the basal hadrosaurid dinosaur Eotrachodon from the uppermost Santonian (Cretaceous) of southern Appalachia. PeerJ, 4, e 1872. doi: 10.7717 / peerj. 1872","Tsogtbaatar, K., Weishampel, D. B., Evans, D. C. & Watabe, M. 2014. A new hadrosauroid (Plesiohadros djadokhtaensis) from the Late Cretaceous Djadokhtan fauna of southern Mongolia. Pp. 108 - 135 in D. A. Eberth & D. C. Evans (eds) Hadrosaurs. Indiana University Press, Bloomington and Indianapolis.","McDonald, A. T., Kirkland, J. I., DeBlieux, D. D., Madsen, S. K., Cavin, J., Milner, A. R. C. & Panzarin, l. 2010 a. New basal iguanodonts from the Cedar Mountain Formation of Utah and the evolution of thumb-spiked dinosaurs. PLoS ONE, 5, e 14075. doi: 10.1371 / journal. pone. 0014075","Xu, X., Tan, Q., Gao, Y., Bao, Z., Yin, Z., Guo, B., Wang, J., Tan, L., Zhang, Y. & Xing, H. 2018. A large-sized basal ankylopollexian from East Asia, shedding light on early biogeographic history of Iguanodontia. Science Bulletin, 63, 556 - 563.","Winkler, D. A., Murry, P. A. & Jacobs, L. L. 1997. A new species of Tenontosaurus (Dinosauria: Ornithopoda) from the Early Cretaceous of Texas. Journal of Vertebrate Paleontology, 17, 330 - 348.","Thomas, D. A. 2015. The cranial anatomy of Tenontosaurus tilletti Ostrom, 1970 (Dinosauria, Ornithopoda). Palaeontologia Electronica, 18 (2), 37 A. doi: 10.26879 / 450","Norman, D. B. 2015. On the history, osteology, and systematic position of the Wealden (Hastings Group) dinosaur Hypselospinus fittoni (Iguanodontia: Styracosterna). Zoological Journal of the Linnean Society, 173, 92 - 189.","McDonald, A. T., Barrett, P. M. & Chapman, S. D. 2010 b. A new basal iguanodont (Dinosauria: Ornithischia) from the Wealden (Lower Cretaceous) of England. Zootaxa, 2569, 1 - 43.","Kirkland, J. I. 1998. A new hadrosaurid from the upper Cedar Mountain Formation (Albian-Cenomanian: Cretaceous) of eastern Utah - the oldest known hadrosaurid (lambeosaurine?). New Mexico Museum of Natural History and Science, Bulletin, 14, 283 - 295.","Kobayashi, Y. & Azuma, Y. 2003. A new iguanodontian (Dinosauria: Ornithopoda) from the Lower Cretaceous Kitadani Formation of Fukui Prefecture, Japan. Journal of Vertebrate Paleontology, 23, 392 - 396.","You, H., Ji, Q. & Li, D. 2005. Lanzhousaurus magnidens gen. et sp. nov. from Gansu Province, China: the largesttoothed herbivorous dinosaur in the world. Geological Bulletin of China, 24, 785 - 794.","Horner, J. R., Weishampel, D. B. & Forster, C. A. 2004. Hadrosauridae. Pp. 436 - 463 in D. B. Weishampel, P. Dodson & H. Osmolska (eds) The Dinosauria. Second edition. University of California Press, Berkeley.","Galton, P. M. 1974. The ornithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight. Bulletin of the British Museum (Natural History), Geology, 25, 1 - 152.","Galton, P. M. 2009. Notes on Neocomian (Lower Cretaceous) ornithopod dinosaurs from England - Hypsilophodon, Valdosaurus, \" Camptosaurus \", \" Iguanodon \" - and referred specimens from Romania and elsewhere. Revue de Paleobiologie, 28, 211 - 273.","Weishampel, D. B., Norman, D. B. & Grigorescu, D. 1993. Telmatosaurus transsylvanicus from the Late Cretaceous of Romania: the most basal hadrosaurid dinosaur. Palaeontology, 36, 361 - 385.","Galton, P. M. 1983. The cranial anatomy of Dryosaurus, a hypsilophodontid dinosaur from the Upper Jurassic of North America and East Africa, with a review of hypsilophodontids from the Upper Jurassic of North America. Geologica et Palaeontologica, 17, 207 - 243.","Prieto-Marquez, A., Chiappe, L. M. & Joshi, S. H. 2012. The lambeosaurine dinosaur Magnapaulia laticaudus from the Late Cretaceous of Baja, California, northwestern Mexico. PLoS ONE, 7, e 38207. doi: 10.1371 / journal. pone. 0038207","Bertozzo, F., Della Vecchia, F. M. & Fabbri, M. 2017. The Venice specimen of Ouranosaurus nigeriensis (Dinosauria, Ornithopoda). PeerJ, 5, e 3403. doi: 10.7717 / peerj. 3403","Carpenter, K. & Wilson, Y. 2008. A new species of Camptosaurus (Ornithopoda: Dinosauria) from the Morrison Formation (Upper Jurassic) of Dinosaur National Monument, Utah, and a biomechanical analysis of its forelimb. Annals of Carnegie Museum, 76, 227 - 263.","Norman, D. B. 2010. A taxonomy of iguanodontians (Dinosauria: Ornithopoda) from the lower Wealden Group (Cretaceous: Valanginian) of southern England. Zootaxa, 2489, 47 - 66.","Gilmore, C. W. 1933. On the dinosaurian fauna of the Iren Dabasu Formation. Bulletin of the American Museum of Natural History, 67, 23 - 78.","Gasulla, J. M., Escaso, F., Narvaez, I., Otega, F. & Sanz, J. L. 2015. A new sail-backed styrocasternan (Dinosauria: Ornithopoda) from the Early Cretaceous of Morella, Spain. PLoS ONE, 10, e 0144167. doi: 10.1371 / journal. pone. 0144167","Romer, A. S. 1956. Osteology of the reptiles. University of Chicago Press, Chicago, 772 pp.","Norman, D. B. 2011. On the osteology of the lower Wealden (Valanginian) ornithopod Barilium dawsoni (Iguanodontia: Styracosterna). Special Papers in Palaeontology, 86, 165 - 194.","Gilpin, D., DiCroce, T. & Carpenter, K. 2007. A possible new basal hadrosaur from the Lower Cretaceous Cedar Mountain Formation of eastern Utah. Pp. 79 - 89 in K. Carpenter (ed.) Horns and beaks: ceratopsian and ornithopod dinosaurs. Indiana University Press, Bloomington and Indianapolis.","Barrett, P. M. & Bonsor, J. A. 2021. A revision of the nonavian dinosaurs Eucercosaurus tanyspondylus and Syngonosaurus macrocercus from the Cambridge Greensand, UK. Cretaceous Research, 118, 1 - 13.","McDonald, A. T. 2011. The taxonomy of species assigned to Camptosaurus (Dinosauria: Ornithopoda). Zootaxa, 2783, 52 - 68.","McDonald, A. T., Wolfe, D. G. & Kirkland, J. I. 2010 c. A new basal hadrosauroid (Dinosauria: Ornithopoda) from the Turonian of New Mexico. Journal of Vertebrate Paleontology, 30, 799 - 812.","Verdu, F. J., Godefroit, P., Royo-Torres, R., Cobos, A. & Alcala, L. 2017 a. Individual variation in the postcranial skeleton of the Early Cretaceous Iguanodon bernissartensis (Dinosauria: Ornithopoda). Cretaceous Research, 74, 65 - 86.","Forster, C. A. 1990. The postcranial skeleton of the ornithopod dinosaur Tenontosaurus tilletti. Journal of Vertebrate Paleontology, 10, 273 - 294.","Brett-Surman, M. K. & Wagner, J. R. 2007. Discussion of character analysis of the appendicular anatomy in Campanian and Maastrichtian North American hadrosaurids - variation and ontogeny. Pp. 135 - 169 in K. Carpenter (ed.) Horns and beaks: ceratopsian and ornithopod dinosaurs. Indiana University Press, Bloomington and Indianapolis.","Ostrom, J. H. 1970. Stratigraphy and paleontology of the Cloverly Formation (Lower Cretaceous) of the Bighorn Basin area, Wyoming and Montana. Bulletin of the Peabody Museum of Natural History, 35, 1 - 234.","Gilmore, C. W. 1925. Osteology of ornithopodous dinosaurs from the Dinosaur National Monument, Utah, Camptosaurus medius, Dryosaurus altus, Laosaurus gracilis. Memoirs of the Carnegie Museum, 10, 385 - 409.","Chanthasit, P. 2010. The ornithopod dinosaur Rhabdodon from the Late Cretaceous of France: anatomy, systematics and paleobiology. Unpublished PhD thesis, Universite Claude Bernard - Lyon, 196 pp. https: // tel. archivesouvertes. fr / tel- 00841228 / file / TH 2010 _ Chanthasit _ Phornphen. pdf","Hooley, R. W. 1925. On the skeleton of Iguanodon atherfieldensis sp. nov., from the Wealden Shales of Atherfield (Isle of Wight). Quarterly Journal of the Geological Society of London, 81, 1 - 61.","Hulke, J. W. 1882. Description of some Iguanodon remains indicating a new species I. seelyi. Quarterly Journal of the Geological Society of London, 150, 135 - 144.","Verdu, F. J., Royo-Torres, R., Cobos, A. & Alcala, L. 2017 b. New systematic and phylogenetic data about the early Barremian Iguanodon galvensis (Ornithopoda: Iguanodontoidea) from Spain. Historical Biology, 30, 437 - 474.","Verdu, F. J., Royo-Torres, R., Cobos, A. & Alcala, L. 2015. Perinates of a new species of Iguanodon (Ornithischia: Ornithopoda) from the lower Barremian of Galve (Teruel, Spain). Cretaceous Research, 56, 250 - 264.","McDonald, A. T. 2012 a. The status of Dollodon and other basal iguanodonts (Dinosauria: Ornithischia) from the upper Wealden beds (Lower Cretaceous) of Europe. Cretaceous Research, 33, 1 - 6."]}

Additional figures and references that did not make it into the final manuscript but still show valuable results.

The geological and paleoenvironmental setting and the vertebrate taxonomy of the fossiliferous, Cenomanian-age deltaic sediments in eastern Morocco, generally referred to as the “Kem Kem beds”, are reviewed. These strata are recognized here as the Kem Kem Group, which is composed of the lower Gara Sbaa and upper Douira formations. Both formations have yielded a similar fossil vertebrate assemblage of predominantly isolated elements pertaining to cartilaginous and bony fishes, turtles, crocodyliforms, pterosaurs, and dinosaurs, as well as invertebrate, plant, and trace fossils. These fossils, now in collections around the world, are reviewed and tabulated. The Kem Kem vertebrate fauna is biased toward large-bodied carnivores including at least four large-bodied non-avian theropods (an abelisaurid, Spinosaurus, Carcharodontosaurus, and Deltadromeus), several large-bodied pterosaurs, and several large crocodyliforms. No comparable modern terrestrial ecosystem exists with similar bias toward large-bodied carnivores. The Kem Kem vertebrate assemblage, currently the best documented association just prior to the onset of the Cenomanian-Turonian marine transgression, captures the taxonomic diversity of a widespread northern African fauna better than any other contemporary assemblage from elsewhere in Africa.

The Solnhofen pterosaurs Pterodactylus antiquus, Aerodactylusscolopaciceps, Diopecephalus kochi, Germanodactylus cristatus and Germanodactylus rhamphastinus all have complicated taxonomic histories. Species originally placed in the genus Pterodactylus, such as Aerodactylus scolopaciceps, Ardeadactylus longicollum, Cycnorhamphus suevicus and Germanodactylus cristatus possess apomorphies not observed in the type species of Pterodactylus, and consequently have been placed in new genera. The affinities of another Solnhofen pterosaur previously placed in Pterodactylus, Diopecephalus kochi, are less clear. It has been proposed that D. kochi is a juvenile specimen of Pterodactylus antiquus, or perhaps “Germanodactylus rhamphastinus” specimens are mature examples of D. kochi. Furthermore, studies have suggested that “Germanodactylus rhamphastinus” is not congeneric with the type species of Germanodactylus. Geometric morphometric analysis of prepubes and a cladistic analysis of the Pterosauria elucidate plesiomorphic and apomorphic conditions for basal Jurassic pterodactyloids. Germanodactylus is found to be a monotypic genus and Pterodactylus, Diopecephalus, and “G. rhamphastinus” are found as distinct taxa belonging in individual genera, diagnosable using a combination of characters. Thus, Diopecephalus kochi is not demonstrated to be congeneric with Germanodactylus or Pterodactylus and is maintained as a valid taxon. “G. rhamphastinus” is readily distinguishable from other Solnhofen pterosaur taxa, and a new genus is erected for its reception.

The taxonomy of the Late Jurassic pterodactyloid pterosaur Pterodactylus scolopaciceps Meyer, 1860 from the Solnhofen Limestone Formation of Bavaria, Germany is reviewed. Its nomenclatural history is long and complex, having been synonymised with both P. kochi (Wagner, 1837), and P. antiquus (Sömmerring, 1812). The majority of pterosaur species from the Solnhofen Limestone, including P. scolopaciceps are represented by juveniles. Consequently, specimens can appear remarkably similar due to juvenile characteristics detracting from taxonomic differences that are exaggerated in later ontogeny. Previous morphological and morphometric analyses have failed to separate species or even genera due to this problem, and as a result many species have been subsumed into a single taxon. A hypodigm for P. scolopaciceps, comprising of the holotype (BSP AS V 29 a/b) and material Broili referred to the taxon is described. P. scolopaciceps is found to be a valid taxon, but placement within Pterodactylus is inappropriate. Consequently, the new genus Aerodactylus is erected to accommodate it. Aerodactylus can be diagnosed on account of a unique suite of characters including jaws containing 16 teeth per-jaw, per-side, which are more sparsely distributed caudally and terminate rostral to the nasoantorbital fenestra; dorsal surface of the skull is subtly depressed rostral of the cranial table; rostrum very elongate (RI = ∼7), terminating in a point; orbits correspondingly low and elongate; elongate cervical vertebrae (approximately three times the length of their width); wing-metacarpal elongate, but still shorter than the ulna and first wing-phalanx; and pteroid approximately 65% of the total length of the ulna, straight and extremely thin (less than one third the width of the ulna). A cladistic analysis demonstrates that Aerodactylus is distinct from Pterodactylus, but close to Cycnorhamphus Seeley, 1870, Ardeadactylus Bennett, 2013a and Aurorazhdarcho Frey, Meyer and Tischlinger, 2011, consequently we erect the inclusive taxon Aurorazhdarchidae for their reception.

The first records of pterosaurs from the Cretaceous of Poland are reported, on the basis of fragmentary remains from the marine Upper Albian (Lower Cretaceous) of the Annopol Anticline, central Poland. The new material consists of four bone fragments, tentatively interpreted as: 1) a portion of wing phalanx; 2) a medial element of fused skull bones (parietal crest?); 3) a fragmentary carpal or tarsal; and 4) a distal phalanx of the pes (or a very small fragment of a long cervical vertebra). Previously, only the remains of marine vertebrates have been reported from the Cretaceous of the Annopol area. The pterosaur fossils studied most probably belonged to individuals that died while over the sea. The possibility that they represent remains dropped from floating carcasses, introduced into the marine environment by rivers, is regarded as less probable, as there are no remains of dinosaurs or other terrestrial fauna in the Annopol deposits.