taxonID	type	description	language	source
03CF5731132B957EFF52871FFC1D1071.taxon	description	Snout shape. The rounded anterior portion of premaxillae and dentaries is a diagnostic character of Lonchodraco giganteus according to Rodrigues and Kellner (2013). In Lonchodraco sp. from the Cenomanian Melovatka Formation of Volgograd Region, Russia, the snout is pointed (Averianov and Kurochkin 2010: fig. 1). The snout shape is unknown in other lonchodectid taxa. This character could be an autapomorphy of L. giganteus. Dorsoventrally flattened jaw tips. This is a diagnostic character of the Lonchodectidae, according to Unwin (2001). This character is present in L. giganteus (Rodrigues and Kellner 2013). In L. compressirostris, the tip of the rostrum likely was transversely compressed rather than dorsoventrally flattened. In Ikrandraco avatar, only the tip of the rostrum is dorsoventrally flattened. In Ikrandraco machaerorhynchus the tip of rostrum is unknown, but the tip of the mandible is flattened dorsoventrally. The taxonomic value of this character is not clear. Alveolar margins divergence. Divergent alveolar margins of the anterior end of the upper and lower jaws is a diagnostic character of Lonchodraco giganteus (Rodrigues and Kellner 2013). According to these authors, L. machaerorhynchus differs from L. giganteus by “ straight ” alveolar margins in dorsal view (Rodrigues and Kellner 2013: 28). “ Parallel margins ” is a more appropriate term in this case as diverging margins are also straight. This term was used in the original description of Ornithocheirus machaerorhynchus by Seeley (1870: 114). The state of this character is unknown for Ikrandraco avatar. Palatal ridge. Longitudinal ridge on the palate was used for diagnosing the genus Lonchodectes (Hooley 1914). Unwin (2003: 179) formulated this character in the diagnosis of Lonchodectes as “ a prominent, sharply ridged, median keel on the occlusal surface of the rostrum. ” A deep palatal ridge is a diagnostic character of Lonchodraco, according to Rodrigues and Kellner (2013). The palatal ridge was indicated for L. giganteus (Rodrigues and Kellner 2013: fig. 4 D) although it is obscured by matrix. A deep palatal ridge was also cited as an autapomorphy of Lonchodraco (?) microdon by Rodrigues and Kellner (2013). This species is considered here to be a synonym of L. machaerorhynchus. In Lonchodectes compressirostris, the palatal ridge is confined to the posterior part of the palate apparently because of the great transverse narrowness of the anterior part of the snout. The palatal ridge is here considered to be a diagnostic character for the Lonchodectidae, while a short posteriorly located palatal ridge is diagnostic for Lonchodectes compressirostris. Palatal surface shape. Palate between the elevation of the alveolar margins and the palatal ridge concave is a diagnostic character of Lonchodraco (?) microdon according to Rodrigues and Kellner (2013). However, the palatal surface is obscured by matrix in L. giganteus. In Lonchodectes compressirostris, the palatal surface between the palatal ridge and alveolar margins has a similar concave shape (Owen 1851 a: pl. 28, fig. 9). This character does not help to distinguish taxa within the Lonchodectidae. Premaxillae dorsal margin. Rounded dorsal margin of premaxillae is a diagnostic character of Lonchodraco (?) microdon, according to Rodrigues and Kellner (2013). This character is redundant because it is correlative with the absence of a premaxillary crest in this taxon. Premaxillary crest. The premaxillary crest is a diagnostic character of Lonchodraco giganteus, according to Rodrigues and Kellner (2013). The absence of a premaxillary crest is a diagnostic character of Lonchodraco (?) microdon, according to Rodrigues and Kellner (2013). According to Rigal et al. (2017), there is no evidence of a premaxillary crest in L. giganteus. In the absence of a definition what constitutes a premaxillary crest it is a matter of opinion to describe the condition seen in L. giganteus as a premaxillary crest or tall premaxilla. In contrast with the mandibular crest, the premaxillary crest has no evident transverse constriction in relation to the rest of the premaxilla. The premaxillary crest is also absent in Lonchodectes and Ikrandraco. Here the premaxillary crest is considered a diagnostic character for the genus Lonchodraco. Odontoid process. Rigal et al. (2017) diagnosed Lonchodraco as having a triangular dorsally directed perforate process at the symphysis extending two to three millimeters above the dental border. This structure corresponds to an odontoid process present in some pterodactyloids (Martill 2014; Kellner et al. 2019 a; Pêgas et al. 2019). As this character has a wider distribution, it cannot be diagnostic for Lonchodraco. Mandibular groove. A deep, V-shaped median sulcus on the occlusal surface of the mandibular symphysis is a diagnostic feature of Lonchodectes according to Unwin (2003). Wide mandibular groove is an autapomorphy of Lonchodraco machaerorhynchus, according to Rodrigues and Kellner (2013). A deep mandibular groove is present in Lonchodraco giganteus, but its details are obscured by matrix (Rodrigues and Kellner 2013). A distinctly wider mandibular groove is present in the anterior mandible fragment NHMUK R 2269 from the Cambridge Greensand, referred to Lonchodectes microdon by Unwin (2001: fig. 11 F) and not mentioned by Rodrigues and Kellner (2013). The mandible is unknown for Lonchodectes compressirostris, but this taxon might not have the mandibular groove because of a strongly transversely narrow snout and short palatal ridge confined to the posterior part of the palate. In Ikrandraco avatar the presence of a mandibular groove cannot be established because of the flattened preservation. This character is a possible synapomorphy for the Lonchodectidae. Mandibular crest. The mandibular crest is a diagnostic character of Lonchodraco, according to Rodrigues and Kellner (2013). Short, low, blade-like dentary crest is an autapomorphy of Lonchodraco giganteus (Rodrigues and Kellner 2013). Deep dentary crest is an autapomorphy of Lonchodraco machaerorhynchus (Rodrigues and Kellner 2013). However, according to my observation, there is no significant difference in the relative dorsoventral depth of the mandibular crest between these two taxa. Rodrigues and Kellner (2013) also noted that L. giganteus differs from anhanguerids in that the mandibular crest does not start at the tip of the mandible. Indeed, the dorsoventrally low mandibular crest in L. giganteus starts some distance posterior to the anterior margin of the mandible and does not increase in depth for some distance. A similar low mandibular crest in the anterior portion of the mandible is present in Lonchodraco microdon (NHMUK R 2269). Rodrigues and Kellner (2013) considered Lonchodectes compressirostris to lack the mandibular crest based on the erroneous interpretation of the anterior rostrum fragment as a fragment of the mandibular symphysis. A mandibular crest is currently unknown for that taxon. A very prominent mandibular crest is present in Ikrandraco. The mandibular crest is presently considered as a diagnostic character for the Lonchodectidae (unknown in Lonchodectes). Ventral margin of mandible posterior to mandibular crest. The ventral margin of mandible posterior to the mandibular crest ascending in lateral view is an autapomorphy of Lonchodraco machaerorhynchus, according to Rodrigues and Kellner (2013). See the next character for discussion. Ventral depression of mandible. Ventral depression located posterior to the mandibular crest is an autapomorphy of Lonchodraco machaerorhynchus according to Rodrigues and Kellner (2013). Seeley (1870) considered a peculiar posterior margin of the mandibular crest of the holotype of Ornithocheirus machaerorhynchus (CAMSM B 54855) as the angular facet, a feature differentiating this taxon from the other pterodactyloids known at that time. In Pteranodon, the angular comes close to the mandibular symphysis but contacts the dentary dorsally, not posteriorly (Bennett 2001: fig. 22 B). Rodrigues and Kellner (2013) interpreted this margin as the posterior surface of the mandibular crest. They proposed two autapomorphies for this taxon based on this interpretation: ventral depression located posteriorly to the dentary crest and ventral margin of the mandible posterior to the dental crest ascending in lateral view. However, this surface is likely the broken surface, and the mandibular crest was continuing posteriorly. This is supported by the ventral margin of the mandibular crest, which deepens posteriorly and probably attained the maximum depth in the missing posterior part. Alveolar parapet. Alveoli placed in an elevation in relation to the palate and the dorsal margin of the mandible is a diagnostic character of the Lonchodectidae according to Unwin (2001, 2003), or an autapomorphy of Lonchodraco, according to Rodrigues and Kellner (2013). This character is present in Lonchodectes compressirostris (contra Rodrigues and Kellner 2013). Here it is considered a diagnostic character of the Lonchodectidae. Raised margins of alveoli. Unwin (2001) considered dental alveoli with margins that are raised into a low collar so that the teeth appear to be “ pedicellate, ” as a character diagnostic for the Lonchodectidae. Later he formulated this character as “ alveoli with margins raised into a low collar ” (Unwin 2003: 179). According to Rigal et al. (2017), the teeth set in raised alveoli, separated by a C-shaped depression, is a diagnostic character for Lonchodraco, a view, which is followed here. Size of alveoli. Comparatively small alveoli (up to 4 mm in diameter) in the anterior portions of the upper and lower jaws is a diagnostic character of the Lonchodectidae or Lonchodraco, according to Unwin (2001, 2003) and Rodrigues and Kellner (2013), respectively. Small alveoli are present also in Lonchodectes compressirostris and Ikrandraco avatar. This character is considered here diagnostic for the Lonchodectidae. Size variation of alveoli. Teeth “ uniform in size ” was cited in the original diagnosis of Lonchodectes (Hooley 1914: 535). “ Subequal sized ” alveoli are diagnostic for Lonchodectes, according to Unwin (2003: 179). Alveoli of the anterior portions of the upper and lower jaws without significant variation in size is a diagnostic character of Lonchodraco according to Rodrigues and Kellner (2013). The teeth do not vary in size in Ikrandraco avatar. The same condition was likely present in Lonchodectes compressirostris, although the tip of the snout is unknown in this taxon. The size variation of alveoli is considered here diagnostic for the Lonchodectidae. Spacing between alveoli. Spacing between alveoli roughly equivalent to their diameters is a diagnostic character of Lonchodraco according to Rodrigues and Kellner (2013). Spacing between alveoli larger than their diameters is an autapomorphy of Lonchodraco (?) microdon according to Rodrigues and Kellner (2013). The latter character contradicts the generic diagnosis of Lonchodraco. In Ikrandraco avatar and I. machaerorhynchus, the spaces between the alveoli are of variable size, some are similar in length with the alveoli, whereas others are larger. Here this character is considered diagnostic for the genus Lonchodraco. The number of alveoli per 3 cm. Approximately six alveoli per 3 cm of jaw margin is an autapomorphy of Lonchodraco giganteus, according to Rodrigues and Kellner (2013). About 4.5 alveoli per 3 cm of jaw margin is an autapomorphy of Lonchodraco machaerorhynchus and, at the same time, is a diagnostic character for Lonchodraco (?) microdon, according to Rodrigues and Kellner (2013). In Ikrandraco avatar, the average number of alveoli per 3 cm is 3.8 for dentary. Rigal et al. (2017) considered tooth count per 3 cm a not taxonomically reliable character because it varies ontogenetically. This could be well true, but the size difference between the holotypes of L. giganteus and I. machaerorhynchus is not significant, and both likely belong to adult individuals. However, the spacing of teeth is markedly different between these two taxa, being denser in L. giganteus. Here these characters are considered diagnostic for the genera Lonchodraco and Ikrandraco. Tooth shape. “ More or less laterally compressed teeth ” is a diagnostic feature of the genus Lonchodectes according to Hooley (1914: 535). According to Unwin (2003), in Lonchodectes, the teeth have constricted bases. Rigal et al. (2017) diagnosed Lonchodraco as having short, conical teeth that are gently recurved labially. However, in most of the lonchodectid specimens, the teeth are not preserved, and the diagnostic value of these characters cannot be evaluated.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132D957EFCDB85EBFC0615A1.taxon	type_taxon	Type genus. Lonchodectes Hooley, 1914.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132D957EFCDB85EBFC0615A1.taxon	diagnosis	Diagnosis. Differs from other pterodactyloid pterosaurs by the combination of the following characters: the palate has a midline ridge; the alveolar margins of upper and lower jaws are elevated in relation to the palatal or dorsal mandibular surface; in the anterior part of the upper and lower jaws, the alveoli are small (up to 4 mm in diameter), without a significant variation in size; mandible with a prominent mandibular crest (unknown for Lonchodectes). Included genera. The type genus, Lonchodraco Rodrigues et Kellner, 2013 and Ikrandraco Wang et al., 2014.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132D957EFCDB85EBFC0615A1.taxon	discussion	Comments. Witton et al. (2009) erroneously cited Hooley (1914) as the author of the family Lonchodectidae.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132D957FFCD880A0FE101174.taxon	type_taxon	Type species. Pterodactylus compressirostris Owen, 1851 (Kuhn 1967: 46).	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132D957FFCD880A0FE101174.taxon	diagnosis	Diagnosis. Differs from Lonchodraco Rodrigues et Kellner, 2013 by low rostrum lacking the premaxillary crest and considerably constricted transversely, and short palatal ridge, confined to the posterior part of the palate. Included species. Type species only.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132D957FFCD880A0FE101174.taxon	discussion	Comments. Lonchodectes cannot be currently distinguished from Ikrandraco Wang et al., 2014 (see comments to that taxon).	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132C957CFF528474FED811AB.taxon	description	(Fig. 1)	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132C957CFF528474FED811AB.taxon	materials_examined	Holotype. NHMUK PV 39410, partial rostrum in two fragments. Type locality and horizon. Culand Pits, Burham, Kent, England; Chalk Formation (Cenomanian-Turonian). Referred specimens. CAMS B 54.584, rostrum fragment; Cambridge, Cambridgeshire, England; Cambridge Greensand (Albian).	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132C957CFF528474FED811AB.taxon	diagnosis	Diagnosis. As for the genus.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132C957CFF528474FED811AB.taxon	discussion	Comments. Two parts of the holotype were considered as parts of a single rostrum in the original description (Owen 1851 a, b) and most subsequent publications. Kellner (1990) challenged this view, considering the smaller part as a fragment of dentary symphysis based on the presence of a medial groove. This view was held in Rodrigues and Kellner (2013). However, what Kellner took for the medial groove is a palatal surface between the two narrowing rostral parapets (Fig. 1 C). The medial groove on lonchodectid dentaries is separated from the alveolar parapet by flat mandibular surface (Fig. 1 C). There is no reason to change the interpretation of NHMUK PV 39410 rostral fragments and no need to renumber the specimens. This taxon might not have the medial groove on the symphysis at all because it has a short palatal ridge placed far behind the supposed symphysis. Following their interpretation of a small part of NHMUK PV 39410 as a mandibular symphysis, Rodrigues and Kellner (2013) consider P. compressirostris to lack the mandibular crest and did not include this taxon in their family Lonchodraconidae. Owen referred to Pterodactylus compressirostris an associated ulna and radius (NHMUK 49004) and two proximal fragments of first wing phalanx (NHMUK 39411 and 49003; Lydekker (1888: 12 )) from the type locality (Owen 1851 a: pl. 30, figs 4 – 5, pl. 32, fig. 2). However, as these bones were not directly associated with the holotype and there at least two other pterosaur species in the type locality, these specimens are not included in the hypodigm of L. compressirostris.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132F957CFF528306FB2112CA.taxon	description	(Fig. 2)	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132F957CFF528306FB2112CA.taxon	description	Ornithocheirus compressirostris: Benton and Spencer 1995: fig. 8.20 D.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132F957CFF528306FB2112CA.taxon	materials_examined	Type locality and horizon. Culand Pits, Burham, Kent, England; Chalk Formation (Cenomanian-Turonian).	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132F957CFF528306FB2112CA.taxon	diagnosis	Diagnosis. As for the genus.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132F957CFF5284D6FED017DB.taxon	type_taxon	Type species. Pterodactylus giganteus Bowerbank, 1846.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132F957CFF5284D6FED017DB.taxon	diagnosis	Diagnosis. Differs from Lonchodectes Hooley, 1914 and Ikrandraco Wang et al., 2014 by premaxillary crest present, spacing between alveoli roughly equivalent to their diameters, approximately six alveoli per 3 cm of alveolar margin, and teeth set in raised alveoli separated by a C-shaped depression. Additionally differs from Ikrandraco avatar Wang et al., 2014 by anterior end of mandible dorsoventrally flattened, from I. machaerorhynchus (Seeley, 1870) by slightly diverging alveolar margins. Included species. Type species and, tentatively, Lonchodraco (?) sp.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132F9572FCDB823FFD1F10F7.taxon	type_taxon	Type species. Ikrandraco avatar Wang et al., 2014.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132F9572FCDB823FFD1F10F7.taxon	diagnosis	Diagnosis. Differs from Lonchodraco Rodrigues et Kellner, 2013 by lack of premaxillary crest and less densely spaced teeth (3.8 – 4.5 per 3 cm). Included species. Type species and I. machaerorhynchus (Seeley, 1870) comb. nov.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF5731132F9572FCDB823FFD1F10F7.taxon	discussion	Comments. Ikrandraco is included in the Lonchodectidae because it possesses three characters diagnostic for this group: the palate has a midline ridge and small alveoli in the anterior part of the upper and lower jaws (up to 4 mm in diameter), without a significant variation in size. The presence of an elevated alveolar margin of upper and lower jaws cannot be established for I. avatar because of its flattened preservation. Currently, Ikrandraco cannot be reliably distinguished from Lonchodectes Hooley, 1914. The latter taxon has somewhat more widely spaced teeth (about 3.3 teeth per 3 cm). Another potential difference is a greater transverse flattening of the rostrum in Lonchodectes. This character cannot be evaluated in Ikrandraco because of its flattened preservation. For this reason, it cannot be excluded that Ikrandraco Wang et al., 2014 is a junior subjective synonym of Lonchodectes Hooley, 1914. Both taxa are currently retained as valid because of incomplete knowledge of Lonchodectes. Some characters cited in the original diagnosis of Ikrandraco cannot be checked in other lonchodectid taxa (slightly arched dorsal margin of the skull above the nasoantorbital fenestrae; lateral depression on the nasal; strongly inclined quadrate (150 °); median hook-like process on the posterior edge of the mandibular crest; two well-developed pneumatic foramina piercing the lateral surface of the axis; a ventral pneumatic foramen on the proximal portion of the second and third wing phalanges). At least some of these characters could be diagnostic for the Lonchodectidae. A very low skull is likely present also in Lonchodectes compressirostris and Ikrandraco machaerorhynchus. These two taxa also share with I. avatar the lack of the premaxillary crest. A deep mandibular crest is also present in Lonchodraco giganteus and I. machaerorhynchus. The humerus of the holotype of I. avatar was described as having “ a warped deltopectoral crest, with the proximal margin rounded ” (Wang et al. 2014: 3). However, the rounded margin of the deltopectoral crest is anterior, not proximal. This crest is undoubtedly not “ warped ” as in ornithocheirid pterosaurs. In the original description, Ikrandraco was referred to as Pteranodontia incertae sedis (Wang et al. 2014). The phylogenetic analysis performed in that paper does not include any lonchodectid taxon. In the analysis by Zhou et al. (2019), Ikrandraco avatar is the sister taxon to Lonchodraco giganteus.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF573113219573FCDB86FFFB821108.taxon	description	(Figs 3, 4) Ptenodactylus machaerorhynchus: Seeley 1869: xvi. Ptenodactylus oweni: Seeley 1869: xvi. Ptenodactylus microdon: Seeley 1869: xvi.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF573113219573FCDB86FFFB821108.taxon	materials_examined	Holotype. CAMSM B 54.855, partial mandibular symphysis. Type locality and horizon. Cambridge, Cambridgeshire, England; Cambridge Greensand (Albian). Holotype. IVPP V 18199, partial skeleton. Type locality and horizon. Lamadong, Jianchang County, Liaoning Province, China; Jiufotang Formation (Aptian). Referred specimens. IVPP V 18406, partial skeleton; Sihedang, Lingyuan County, Liaoning Province, China; Jiufotang Formation (Aptian). The third specimen of I. avatar was announced by Chen et al. (2018).	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF573113219573FCDB86FFFB821108.taxon	diagnosis	Diagnosis. Differs from I. machaerorhynchus by a relatively deeper mandibular crest and approximately 3.8 alveoli per 3 cm of alveolar margin.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF573113219573FCDB86FFFB821108.taxon	discussion	Comments. In the supplementary information to Wang et al. (2014), a specimen number IVPP V 18904 was applied to both the holotype and the referred specimen. large mandibular crest. Ornithocheirus machaerorhynchus was previously reconstructed as having the deepest point of the mandibular crest near the anterior end of the dentary (Unwin 2001: fig. 12 D), but there is no evidence for this. Here it is reconstructed as having the deepest point of the mandibular crest at the middle of the mandibular symphysis (Fig. 4 D), similar to the condition present in L. giganteus and I. avatar.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF573113209576FC508491FD3D116B.taxon	description	BEXHM 2015.18. It is a fragmentary pterosaur skeleton from the Upper Tunbridge Wells Sand Formation or Lower Weald Clay Formation (upper Valanginian or lower Hauterivian) at Foreshore at Bexhill, East Sussex, England. It includes a jaw fragment with three teeth, six articulated dorsal vertebrae, distal ulna fragment, proximal syncarpal, and fragments of wing phalanges (Rigal et al. 2017). The jaw is likely part of the mandibular ramus posterior to the symphysis. The teeth are small with conical crowns, evenly spaced and located in alveoli with prominent, dorsally raised borders. The dorsal vertebrae are not fused. According to the original description, the anterior-most complete vertebra has an articular facet for the rib that is significantly larger than that of all of the more posterior vertebrae. Rigal et al. (2017) think that the enlarged size of the parapophysis of the anterior-most complete vertebra suggests that it is a free dorsal vertebra. I assume that the authors mean the height of the transverse process rather than the size of the rib articulation facet. The rib articular surfaces (parapophysis and diapophysis) appear to be missing in all vertebrae. The preserved cross-section of the transverse process of the first vertebra is not significantly larger than in other vertebrae. The free vertebrae have a high transverse process but do not have an enlarged parapophyses (articular facet for the rib capitulum). They have a joined articular facet for a single-headed rib, and this facet is distinctly smaller than rib articular facets in anterior dorsals. In free dorsals, the height of the transverse process is more or less uniform. In BEXHM 2015.18, the transverse process of the first complete dorsal vertebra is, however, higher than in other vertebrae. In the two succeeding vertebrae, the height of the transverse process decreases and then slightly increases in the more posterior vertebrae. In Pteranodon, the dorsal vertebrae with a similarly high transverse process is the fifth dorsal, which is followed by dorsals with the decreased height of the transverse process (Bennett 2001: fig. 46). The fifth and sixth dorsals are incorporated in the notarium in Pteranodon. The lower neural spines of dorsal vertebrae in BEXHM 2015.18 also suggest that they are anterior dorsals, not posterior dorsals, as was assumed by the Rigal et al. (2017). It is either this specimen is immature with unfused notarium, and missing supraneural plate or notarium was shorter in this taxon. Rigal et al. (2017) identified in BEXHM 2015.18 a distal fragment of the left ulna and right proximal syncarpal. However, the ulna is the right one according to its morphology. It is more likely to find the association of the bones from a wing of the same side, not a mixture of right and left wing elements, especially for the distal ulna and proximal syncarpal that are articulated in life and closely approximated postmortem in BEXHM 2015.18. The exposed flattened surface of the ulna indeed accommodated the radius, but this is the anterior ulna surface, not posterior surface, as indicated in Rigal et al. (2017: fig. 4 b). The proximal syncarpal retains a suture between its ulnar and radial parts. This is in line with likely not ossified notarium in BEXHM 2015.18 and suggests a juvenile age for this specimen. Rigal et al. (2017) also indicated fragments of the second, third, and fourth phalanges of the left wing digit. However, these fragments are poorly preserved and do not preserve any characters that may suggest an attribution to the left or right side wing. As was noted above, all wing elements in BEXHM 2015.18 appear to be from the right side. Also, I cannot follow the attribution of these fragments to the second, third, and fourth phalanges. Some of these fragments could be pieces of the first wing phalanx. The idea that a subtriangular cross-section of the fourth wing phalanx may be a diagnostic character for the Lonchodectidae is unfounded because identification of this piece of bone as a fourth wing phalanx is dubious and attribution of BEXHM 2015.18 to the Lonchodectidae is poorly supported. BEXHM 2015.18 was referred to the Lonchodectidae because of flattened, simple cone-shaped tooth crowns in alveoli with raised borders (Rigal et al. 2017). The latter character was considered an autapomorphy of the Lonchodectidae by Unwin (2001). Rigal et al. (2017) differentiated BEXHM 2015.18 from Serradraco sagittirostris by having vertically directed teeth and raised alveolae with symmetric borders. According to these authors, BEXHM 2015.18 agrees well in these features with Lonchodraco giganteus and for this reason this specimen was referred to Lonchodraco sp. However, the tooth inclination may vary within the jaw, with more vertical teeth anteriorly. The only known specimen of S. sagittirostris (NHMUK R 1823) comprises posterior dentary fragments. I cannot confirm that the tid Ikrandraco avatar (Wang et al. 2014: fig. 3 e) has a similar plesiomorphic, not “ warped ” deltopectoral crest with parallel margins (contra the original description). Still, this crest is distinctly shorter than in “ Palaeornis. ” The morphology of NHMUK 2353 and 2353 a was reviewed by Averianov (2012), who considered it as belonging to a basal azhdarchoid. The “ Moon Goddess. ” A well pre- served pterosaur skeleton from the Ap- tian Jiufotang Formation of Liaoning Province, China has been attributed to still undescribed lonchodectid taxon, in- formally known as the “ Moon Goddess, ” or “ Chang-e ” (Unwin et al. 2008; Witton 2013). Lonchodectids are present in the Jiufotang Formation, as shown by Ikran- draco avatar.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
03CF573113209576FC508491FD3D116B.taxon	description	SMNK PAL 6597 from the Albian Fig. 5. Dentary fragments BEXHM 2015.18 (A) and NHMUK R 1823, holotype of Serradraco sagittirostris (Owen, 1874) (B) (after Rigal et al., 2017: figs Romualdo Formation in Santana do Cari- 5 a and 7 c). Arrows indicate asymmetric alveolar borders. Scale bar = 10 cm. ri region, Ceará Province, Brazil (Martill 2011). The genus is characterized by teeth in BEXHM 2015.18 have raised alveoli with reduced dentition with only seven tooth pairs antesymmetric borders. This interpretation is based on rior to the nasoantorbital fenestra. The teeth are of a poorly preserved part of the jaw with most of the similar size, in contrast with the heterodont dentition bone is missing. Moreover, the better preserved al- of ornithocheiroid taxa. Originally, the taxon was veolar margin posterior to the preserved teeth shows referred to the Ctenochasmatoidea (Martill 2011). the asymmetric alveolar borders, so characteristic Witton (2013: 211) cited personal communication for S. sagittirostris (Fig. 5). Serradraco sagittirostris from D. M. Unwin that “ several features of its jaw and comes the Upper Tunbridge Wells Sand Formation, dental morphology are consistent with a lonchodectid from where BEXHM 2015.18 may also come. The identity. ” Indeed, the long and low rostrum resembles latter specimen is more likely belong to S. sagitti- that of Lonchodectes and Ikrandraco, and the variarostris and thus excluded from the Lonchodectidae. tion of the tooth size is not significant. Nevertheless, Palaeornis cliftii Mantell, 1844. This species the tooth size variation is more pronounced than in is based on an isolated humerus broken in two parts lonchodectids: the terminal teeth are somewhat big- (NHMUK 2353 and 2353 a) from the Hastings Beds ger and separated from the lateral teeth by a small Group (probably from the Valanginian Upper Tun- tooth. This condition is more reminiscent of the orbridge Wells Formation) at Cuckfield, West Sussex, nithocheiroid dentition. Unwindia could represent an England (Witton et al. 2009: figs 3 – 4). Witton et al. ancestral stage in teeth reduction leading to the Lon- (2009) referred this specimen to the Lonchodectidae chodectidae, but cannot be included in that group. indet. based on its similarity with isolated humerus Prejanopterus curvirostris Fuentes Vidarte et CAMSM B 54.081 from Cambridge Greensand, Meijide Calvo, 2010. Prejanopterus is known from which Unwin (2003) referred to Lonchodectes sp. disarticulated cranial and postcranial elements of However, there are no grounds for referral of the several individuals from the Aptian Leza Formation latter humerus to Lonchodectes, and reference of at Fuente Amarga, Préjano, La Rioja Province, Spain the “ Palaeornis ” humerus to the Lonchodectidae is (Fuentes Vidarte and Meijide Calvo 2010). Prejansimilarly groundless. The humerus of the lonchodec- opterus was originally referred to Pterodactyloidea incertae sedis, but then assigned to the Pterodactylidae (Pereda Suberbiola et al. 2012). According to Witton (2013), “ the long, low nature of its jaw and unusual tooth socket morphology tentatively suggest it might represent a lonchodectid. ” Similarities with the Lonchodectidae include a low elongated rostrum lacking the premaxillary crest (present in Lonchodraco but not in Lonchodectes and Ikrandraco) and small homodont teeth. However, Prejanopterus lacks raised alveolar margins, palatal ridge, and mandibular crest. These characters do not allow the attribution of Prejanopterus to the Lonchodectidae.	en	Averianov, A. O. (2020): Taxonomy of the Lonchodectidae (Pterosauria, Pterodactyloidea). Proceedings of the Zoological Institute of the Russian Academy of Sciences 324 (1): 41-55, DOI: 10.31610/trudyzin/2020.324.1.41, URL: https://doi.org/10.31610/trudyzin/2020.324.1.41
