identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
2B04CD60072DFFE9FC2CCC14F0A8FCC0.text	2B04CD60072DFFE9FC2CCC14F0A8FCC0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Diplocynodon Pomel 1847	<div><p>Genus Diplocynodon Pomel, 1847</p><p>REMARK</p><p>After Martin et al. (2014) the following combination of characters is diagnostic for the genus Diplocynodon: 1) the fourth and fifth maxillary alveoli and the third and fourth dentary alveoli are enlarged and confluent; 2) the lacrimal is longer than the prefrontal; 3) the ectopterygoid is situated close to the posteriormost maxillary alveoli; 4) the dorsal margin of the infratemporal fenestra is bordered by the quadratojugal, preventing the quadrate from reaching the fenestra; and 5) 16-17 alveoli are present in the maxillae. Additional synapomorphy present in all members of the genus Diplocynodon (if preserved), identified by Rio et al. (2020), is that the nasals are excluded at least externally from the naris, while another two characters may be considered as ambiguous synapomorphies that are present in all Diplocynodon species, but also in other taxa, as follows: 1) the frontoparietal suture is linear between the supratemporal fenestrae (concavo-convex in D. ratelii Pomel, 1847, but linear also in Paleosuchus); and 2) the parietal and squamosal approach each other on the posterior wall of the supratemporal fenestrae (Rio et al. 2020). A single cranial autapomorphy of Diplocynodon has been identified by Martin et al. (2014): the quadrate-pterygoid suture line is linear from the basisphenoid exposure to the foramen ovale.</p></div>	https://treatment.plazi.org/id/2B04CD60072DFFE9FC2CCC14F0A8FCC0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Venczel, Márton;Codrea, Vlad A.	Venczel, Márton, Codrea, Vlad A. (2022): A new late Eocene alligatoroid crocodyliform from Transylvania. Comptes Rendus Palevol 21 (20): 411-429, DOI: 10.5852/cr-palevol2022v21a20, URL: https://doi.org/10.5852/cr-palevol2022v21a20
2B04CD60072CFFE3FEA2C8D3F364F8BD.text	2B04CD60072CFFE3FEA2C8D3F364F8BD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Diplocynodon kochi Venczel & Codrea 2022	<div><p>Diplocynodon kochi n. sp.</p><p>(Figs 1; 3-7)</p><p>urn:lsid:zoobank.org:act: C2CA2BAD-656C-4765-BB1C-50BE3CDA6A50</p><p>DERIVATION OF NAME. — A tribute to Koch Antal, eminent geologist from Transylvania, Cluj-Napoca (Kolozsvár, Klausenburg), who brought an important contribution to the field of Transylvanian stratigraphy.</p><p>HOLOTYPE. — UBB V.1453, a three-dimensionally preserved, nearly complete skull lacking from its right side the anterolateral margin of premaxilla, posteriormost part of maxilla, anterior part of jugal and from its left side the postorbital, squamosal, jugal and quadratojugal.</p><p>TYPE LOCALITY. — Cluj-Mănăștur former limestone quarry, Cluj-Napoca, Transylvania, Romania (Fig. 2).</p><p>STRATIGRAPHIC HORIZON AND AGE. — The geological strata, from where the new crocodyliform skull originates, belong to the late Eocene (Priabonian) Cluj Limestone Formation (Mészáros 2000). The stratigraphic unit consists of a series of three coarse bioclastic limestone beds rich in calcareous algae, ostracods, gastropods and bivalve shells (e.g. Anomia tenuistriata Deshayes, 1832) alternating with calcareous or clayey marls (Koch 1894), which were accumulated in a shallow marine carbonate platform. Occasionally, this carbonate platform emerged and was exposed to atmospheric erosion (Codrea et al. 1997). After Koch (1894), the main layers from the Cluj-Mănăștur limestone quarry are as follows: 1) coarse grained limestone bank (one metre thick, used as building stones or carved stones) with abundant ostracods, moulds of gastropods (mainly Anomia tenuistriata) and rare vertebrate remains (e.g. “ Delphinus sp. ”); 2) fissured shale marlstone (0.2 meters thick) with ostracods and Anomia; 3) coarse grained limestone (0.5 meters thick) with ostracods and extremely small gastropods used as building stones or carved stones; 4) fissured shale marlstone (one meter thick) with ostracods and Anomia tenuistriata; 5) coarse grained limestone (0.8 meters thick) with ostracods and extremely small gastropods and rare Anomia; it represents the uppermost limestone horizon from the Cluj-Mănăștur limestone quarry; and 6) fissured shale marlstone (one meter thick) with numerous chalk-like white limestone concretions; it represents the uppermost sedimentary layer from the quarry. After Koch (1894), a partial skull of a small crocodilian consisting of “a maxilla and several inner bones of the skull” has been recovered by quarry workers. It somewhat resembled Crocodylus communis (i.e., C. niloticus Laurenti, 1768) but not identical with that because the fossil specimen possessed a higher number of teeth (21-22) vs Crocodylus having a lower tooth count (Koch 1894).</p><p>DIAGNOSIS. — Small-sized late Eocene alligatoroid crocodyliform with estimated total body length up to 1.8 m. Diplocynodon kochi n. sp. is diagnosed within that genus by the following unique combination of characters (autapomorphy marked by *): 1) premaxillary surface with deep notch lateral to naris; 2) incisive foramen larger than half of the greatest width of premaxillae; 3) occlusion pits on the premaxillae positioned lingually to alveoli; 4) preorbital ridge prominent and wide with lateral overhang; 5) dermal bones of skull roof overhang rim of supratemporal fenestra; 6*) medial wall of parietal with deep recess; 7) anterior process of quadratojugal short; 8) margin of otic aperture inset from paroccipital process; 9) lateral carotid foramen opens dorsal to basisphenoid at maturity; and 10) posterior maxillary teeth and alveoli mediolaterally compressed.</p><p>DESCRIPTION OF THE HOLOTYPE</p><p>General description</p><p>The holotype is a three-dimensionally preserved skull of a mature individual with portions of the skull damaged at the time of unearthing. The anterolateral part of the right premaxilla is broken off, whereas on the left side a small portion from the medial premaxillary border is missing; the right posterior maxillary ramus, except its posteriormost part preserving two tooth positions, is broken off; the right ectopterygoid is missing; the posterior half of the palatines and the lateral part of the right pterygoid are broken off. The damages observed on the left dorsolateral side of the skull may be related to taphonomic processes. Apparently these bones may have been lost prior to compaction of the embedding sediment as it follows: from the left postorbital only its anteromedial part in articulation with the frontal is preserved; the left squamosal is missing completely, exposing on the posterodorsal side of the otic recess and dorsal to the cranio-quadrate passage its articular surface with the underlying quadrate; the left quadratojugal and jugal are missing, from the latter only an imprint on the posterolateral side of the maxillary ramus is preserved; the left ectopterygoid is shifted anteriorly from its articulation position with the pterygoid and its anterior ramus is broken off. The outer surface of the skull is covered by a strong sculpture consisting of densely distributed, circular or irregular pits and grooves. The sculpture on the preorbital area and on the posterodorsal surface of the maxillae tends to form more elongated grooves, delimited by irregular crests. In dorsal view, the skull is elongated with a narrow snout; the preorbital length/total skull length ratio is of about 66% (Table 1). The premaxillae enclose completely the roughly circular shaped external nares that are positioned closer to the anterior premaxillary margin than to the level of the premaxillary-maxillary notch (Fig. 3). The premaxillary-maxillary notch is anteroposteriorly wide and mediolaterally shallow and flanked by prominent bony ridges that extend on the posterolateral sides of the premaxillary posterior processes. The rostrum is concave dorsally, whereas the interorbital dorsal surface of the frontal is faintly convex. The skull table is flat and the frontoparietal suture is almost straight and situated roughly at the posterior limit of the shallower anterior portion of the supratemporal fenestra. The orbits are twice larger longitudinally than the supratemporal fenestra. The infratemporal fenestra is triangle-shaped and its posteroventral corner is bounded by the quadratojugal. In lateral view, the skull is platyrostral with the dorsal surface of the rostrum slightly curved upward, however, provided with robust and bulging premaxillary narial rims and with a dorsoventral constriction posterior to the external naris (Figs 4; 5). A wide preorbital ridge arises from the posteromedial part of the lacrimal and extends anterolaterally depicting a lateral curvature. The cranial table is formed by the postorbitals and squamosals and their fusion is situated about half-length of the supratemporal fenestra; the otic aperture is inset from the paroccipital process. The orbit is large with a strongly concave medial margin, bordered medially by the frontal and anteromedially by the prefrontal and the lacrimal. The lateral margin of the orbit is smoothly sinuous with a somewhat elevated anterolateral margin bounded by the lacrimal, whereas a less elevated posterolateral margin is bordered by the dorsally convex anterior ramus of the jugal representing the lower periorbital crest (Andrade &amp; Hornung 2011); the posterior margin of the orbit is separated from the infratemporal fenestra by a pillar-like postorbital bar, resulted from the fusion of the dorsally situated flange of the postorbital with the ventrally situated and strongly inset jugal dorsal process. The infratemporal fenestra is wider and longer than the supratemporal fenestra and delimited posteriorly by the quadratojugal. In ventral view, the suborbital fenestrae are elongated and large with concave medial margins, whereas the lateral margins are almost straight (Fig. 6). Their most anterior limit reaches the level of the tenth alveolus. The suborbital fenestrae are bounded medially and anteriorly by the anteriorly widening palatines, and laterally by the maxillae, whereas posterolaterally by the anterior processes of the ectopterygoids; posteriorly these are bounded by the pterygoids. However, the articulation between the palatines and pterygoids cannot be discerned on the specimen. As a consequence, we may only presume that it was near to the posterior limit of the suborbital fenestrae, as it is seen in other members of Diplocynodon (Rio et al. 2020: fig. 2C, D). The secondary choana opens anteriorly from the posterior border of the pterygoids, however, it is strongly damaged with part of the delimiting bony laminae broken off. Fortunately, the embedding sediment preserves considerable part of the choanal morphology indicating that it has been heart-shaped with its anterior margin flush with the pterygoid surface, whereas its lateral margin is delimited by a slightly depressed area; the choanal septum is recessed within the choana (Fig. 6).</p><p>Premaxilla. The premaxillary rim, enclosing the external nares, is faintly bulging on its posterior half and bordered posterolaterally by a deep notch connected to several pits and transversal grooves; the anterior side of the premaxillary rim is flush with the narial opening. The posterior premaxillary processes extend back to the level of the third maxillary tooth positions. These processes are sutured dorsally and as a consequence the nasals are excluded broadly from the narial margins (Fig. 3).</p><p>In ventral view, the incisive foramen is relatively large (i.e., larger than half of the greatest width of the premaxillae) and oval in shape, situated at some distance from the premaxillary tooth row, where it extends between the levels of the second and fourth alveoli (Fig. 6). There are five alveoli in the premaxillae, of which the third and fourth are the largest. On the ventral side of the left premaxilla, three occlusal pits are discernible lingually to the tooth row: the first pit is larger and placed between the levels of the first and second alveoli; the second pit is smaller, placed posteromedially to the third alveolus; the third pit is small and situated posteromedially to the fifth alveolus. The suture line with the maxillae is faintly convex anteriorly, situated at the posterior border of the premaxillary-maxillary notch.</p><p>Maxilla. The outer surface of the maxillary nasal ramus is convex labially, whereas its anterolateral surface near the premaxillary fusion line is shallowly concave, as it is the posterior maxillary ramus, covered by the jugal (Fig. 3). In dorsal or ventral views, the lateral margin is produced into a lateral convexity between the first and sixth maxillary alveoli and the widest point is reached at the level of the fourth-fifth largest alveoli. The maxillary tooth row most probably included 16 teeth, of which the fourth and fifth are the largest with nearly confluent alveoli of roughly the same size; the posterior maxillary alveoli, starting from the sixth alveolus, are positioned in a straight line. As preserved, the roots or alveoli of the posteriormost teeth (positions 12th-16th) are slightly compressed mediolaterally. The occlusal pits, up to the fifth alveolus, are positioned lingually to the alveoli; the remaining occlusal pits, visible between the fifth-twelfth alveoli, are situated in line with the tooth row; posterior to the 12th maxillary alveolus, there is no trace of occlusal pit. The foramen for the palatine ramus of the fifth cranial nerve is visible on the left side, medially to the eighth maxillary alveolus (Fig. 6).</p><p>Nasal. The nasals are elongated extending from the level of the first to the 12th maxillary tooth positions (Fig. 3). The nasals are restricted anteriorly by the paired premaxillae, whereas posteriorly are sutured with the frontal medially, and with the prefrontal and lacrimal laterally.</p><p>Lacrimal. These bones are well preserved, except the anterior portion of the right lacrimal that is broken off together with the posterior ramus of the maxilla (Fig. 5A). The lacrimal is elongated and extends more anteriorly than the prefrontal, wedged between the maxilla and the nasal. The posterolateral part of the lacrimal is sutured with the jugal, whereas its posteromedial edge encloses the anterolateral margin of the orbit exposing there a smooth, boomerang-shaped surface. The preorbital ridge extends anterolaterally, arising from the smooth, posteromedial part of the lacrimal, depicting a lateral curvature, and it terminates at the level of the anterior limit of the jugal; it possesses a prominent bulging peak with a lateral overhang (Fig. 4A, C).</p><p>Prefrontal. The prefrontal is more or less a lanceolate shaped bone, tapering both anteriorly and posteriorly. Its anterior extent, positioned slightly anterior to the frontal process, is restricted by the posterior border of the nasolacrimal suture. The medial border of the prefrontal is sutured to the frontal process, whereas its posterolateral part delimits the anteromedial margin of the orbit (Fig. 3). On the left dorsolateral side, small part from the prefrontal pillar extending downward to articulate with the dorsolateral margin of the palatine is exposed.</p><p>Jugal. The jugal is an elongate and anteriorly widening bone, which delimits lateroventrally the orbit and the infratemporal fenestra. However, its greatest vertical thickness is reached at the level of its articulation point with the posterior maxillary ramus. The anterior ramus of the jugal covers dorsolaterally the maxilla with its most anterior part tapering and wedged between the maxilla and the lacrimal. Detail of this morphology is exposed on the left maxilla by an imprint of the lost anterior process of the jugal (Fig. 4A, C). The posterior ramus of the jugal, preserved on the right side only, is rod-like and articulates with the quadratojugal; the articulation point is close to the posterior corner of the infratemporal fenestra exposing a short lateral process of the quadratojugal which protrudes onto the medial surface of the jugal (Fig. 5C). On the medial surface of the jugal, two large foramina are situated just anterior to the ascending process of the jugal and at the level of the posterior maxillary terminus respectively. The ascending process is detached medially from the body of the jugal by the dorsal crest and it is inclined dorsomedially (Fig. 3).</p><p>Frontal. The frontal is a relatively long, dagger-like bone reaching its widest point between the postorbitals (Fig. 3). There is no frontal step in the preorbital area. The rostral part is extremely thin extending anteriorly approximately to the level of the most anterior margin of the jugal. The frontal process penetrates between the posterior rami of the nasals, but it remains slightly shorter than the anterior margins of the prefrontals. Posteriorly, the articulation with the parietal is straight and the suture line extends laterally across the anterior elevated part of the supratemporal fenestra, where the parietal contacts the postorbital. The frontal bounds the anteromedial corner of the supratemporal fenestra, and its contact with the postorbital is located in the anterior margin of the supratemporal fenestra. The dorsal surface of the frontal is flat in the posterior half of the interorbital area with a slightly elevated point at the posterior orbital margin, whereas in the anterior half of the interorbital space its surface is anteroposteriorly convex and it becomes shallowly concave on the frontal process.</p><p>Postorbital. The postorbital encloses the anterolateral part of the supratemporal fenestra. It has a short and relatively thin body that is gently curved and slanting anterolaterally (Figs 3; 4A). The pillar-like postorbital bar is compressed mediolaterally and curved ventrolaterally articulating with the dorsal process of the jugal. The dorsal surface is irregular with some signs of erosion; however, the intact surface bears a strong sculpture consisting of irregular ridges and various sized pits; the sculpture extends downwards and covers also the dorsolateral part of the postorbital bar (Fig. 5A).</p><p>Parietal. The parietal bounds medially the supratemporal fenestra. Its rims are smooth, slightly raised and overhanging laterally the supratemporal fenestra, as observed on the left side of the skull (Fig. 4A, C). Below that overhang, the wall of the parietal is steeply bent medially and produced into a deep recess, probably serving as an extended insertion surface for the jaw adductor muscles (i.e., m. adductor mandibulae externus profundus – MAMEP). The parietal – squamosal contact is placed at the posterior corner of the supratemporal fenestra and their irregular suture line extends back towards the occipital margin. Near the posteromedial border, the supraoccipital prevents the parietal from reaching the occipital margin (Fig. 3). On the anterolateral margin of the supratemporal fenestra, the parietal is sutured to the medial edge of the postorbital, as seen in other members of the genus (e.g. D. remensis Martin et al., 2014) (Martin et al. 2014). Anterolaterally from the posterior wall of the left supratemporal fenestra, from where the squamosal is detached from its articulating point, the dorsal surface of the quadrate is invaded by the parietal approaching close to the sutural surface left by the squamosal (Fig. 4A, C), a condition present in almost all members of Diplocynodon (Rio et al. 2020) .</p><p>Squamosal. The squamosal bounds posterolaterally the supratemporal fenestra (Fig. 3). The dorsal surface is flat and wide having a relatively narrow anterior (i.e., postorbital) process. The articulation with the postorbital is about half-length of the supratemporal fenestra where the squamosal underlaps the postorbital. On the lateral margin of the squamosal, starting from the dorsally less elevated posterior prong, a roughly parallel groove for the upper ear lids extends anteriorly. The sutured line with the underlying quadrate extends from the posterolateral surface of the quadrate up to the posterodorsal side of the otic incision. In lateral view, the posterior margin of the otic aperture forms a well-defined oval recess (Fig. 5A, B), contrasting in this respect with some members of Diplocynodon (see Discussion), in which the posterior margin of the otic aperture is continuous with the paroccipital process (Martin et al. 2014).</p><p>Quadrate. The posterior quadrate rami, preserved on both sides, are elongate and moderately widened (Fig. 7A, B). The dorsal surface of the quadrate is smooth and its anterolateral margin, sutured to the quadratojugal, is flattened; anteriorly, the quadrate is restricted by the quadratojugal to enter in the infratemporal fenestra. Medially, a large concavity is divided ventrally by the otic buttress (sensu Montefeltro et al. 2016): the posterior one represents the incision of the otic aperture of the cranioquadrate passage, bounded completely by the quadrate; anterior to the cranioquadrate passage is situated the otic incision, whereas a smaller anterior opening corresponds to the subtympanic foramen (Fig. 5A). On the left side, the quadrate condyles are dorsally eroded, whereas on the right side the medial hemicondyle is broken off. The quadrate condyle consists of a ventrally deflected medial hemicondyle facing posteriorly and ventrolaterally, whereas the lateral hemicondyle has a posteroventral orientation; the foramen aëreum is situated in a shallow notch on the dorsal surface of the quadrate at some distance to the medial margin and anterior to the medial hemicondyle. In ventral view, the scar for the attachment of the adductor mandibulae posterior muscle is exposed on the posteroventral surface of the quadrate in form of a faint ridge (Fig. 6).</p><p>Quadratojugal. The quadratojugal, preserved on the right side only, is a dorsoventrally flattened bone; its dorsal surface, in line with the posterior process of the jugal, is covered by a pit and ridge sculpture resembling that of the jugal (Fig. 5A). Medially, the quadratojugal is sutured to the lateral hemicondyle of the quadrate, whereas its relatively short anterior process is sutured to the medial side of the posterior ramus of the jugal (Fig. 5C). The quadratojugal process extends along the posterior margin of the infratemporal fenestra excluding the quadrate entirely from that margin; there is no trace of a quadratojugal spine (Fig. 5C).</p><p>Palatine. The anterior half of the paired palatines is preserved, whereas the posterior part is present as an imprint only (Fig. 6). The palatines form the medial margins of the suborbital fenestrae; the anterior part flares anteriorly and sends a lateral process entering into the anteromedial corner of the suborbital fenestra. The anterior flange of the palatine reaches to the level of the eighth maxillary alveolus; the suture with the maxilla is irregular and slightly convex anteriorly, the palatine underlying the maxilla at some distance. The broken surface of the palatine reveals that the choanal septum is complete, V-shaped and possesses a ventral vertical lamina that reaches the ventral surface of the palatine (Fig. 6A).</p><p>Pterygoid. The ventral part of the right pterygoid wing is completely broken off; the left wing is partially preserved (Fig. 6). The conserved parts and the imprints of the broken elements left on the embedding sediment suggest that the secondary choana was completely bounded by the pterygoid. The choanal septum is present, consisting of an extremely thin bony lamina that delimits dorsally a moderately deep embayment representing the medial and posterior corner of the secondary choana. The imprint left by the left choana is oval suggesting that the reconstructed choanae may have been heart-shaped, as in the other members of Diplocynodon (Rio et al. 2020) . The posterior processes of the pterygoid are oriented posterodorsally and slightly laterally.</p><p>Ectopterygoid. The only preserved part is the anterior and posterolateral flange of the left ectopterygoid (Figs 4A, C; 6A, B). The latter is shifted anteriorly from its articulating position with the lateral wing of the pterygoid. The anterior extent of the ectopterygoid is preserved in form of an imprint on the posteromedial side of the left maxillary ramus, which demonstrates that it reached only the level of the last three alveoli and bordered only the last alveolus, as seen in some members of Diplocynodon (see below: Comparisons). The sutured surfaces, exposed on the remnant of the dorsal anterior process of the left ectopterygoid (Fig. 4A, C) and those on the medial side of the right jugal and postorbital bar (Fig. 5C), suggest that the ectopterygoid was firmly sutured to the jugal and to the base of the postorbital bar.</p><p>Supraoccipital. The supraoccipital is exposed dorsally on the medial posterior border of the occipital margin in form of a less elevated area (Fig. 3); the post-temporal fenestrae are visible on both sides of this dorsally exposed area, connected to two ventrally extending furrows; on the left side, from where the squamosal is broken off, the supraoccipital is exposed again dorsally (Fig. 7A, B). In posterior view, the supraoccipital appears as a broad triangle wedging between the sutured exoccipitals above the foramen magnum, thus being excluded from the dorsal margin of that foramen; the occipital surface is deeply concave with a low sagittal ridge on the dorsal half of the bone.</p><p>Exoccipital. The exoccipitals bound the dorsal and lateral margins of the foramen magnum (Fig. 7A, B). The lateral sides of both exoccipitals are more or less damaged. The right lateral paroccipital process is better preserved, and sutured to the prong of the squamosal dorsally and to the vertical lamina of the quadrate ventrally. The cranio-quadrate passage is exposed on the left side, enclosed dorsally by the paroccipital process; medially from that point, a prominent, but relatively short crest, that may correspond to the crista tuberalis (Serrano-Martínez et al. 2019), is developed at level with, but not reaching the foramen magnum. The exoccipital bears three foramina in a common recess, as follows: two small foramina, near the foramen magnum represent the exit of the paired hypoglossal nerves (cranial nerve XII); anterolaterally to these foramina, is a larger foramen representing the exit of the glossopharyngeal and vagus nerves (for cranial nerves IX-XI); ventral to this group of foramina, outside the recess enclosing the above foramina and approximately at the ventral level of the occipital condyle, is situated the lateral carotid foramen (Fig. 7C). On the dorsolateral margin of the lateral carotid foramen a sharp crest is developed and extends dorsolaterally to form the metotic crest that represents the contact area between the exoccipital and the quadrate.</p><p>Basioccipital. The basioccipital forms the floor of the foramen magnum and more ventrally it delimits the median eustachian foramen that opens between the basioccipital plate and the basisphenoid. The basioccipital condyle is of small size, whereas the basioccipital plate is slightly wider than the condyle (Fig. 7A, B); the basioccipital plate faces posteriorly and there is a well-defined sagittal crest extending from the base of the condyle down to the ventral tuberosity; near the condyle, on both sides of the sagittal crest, the bone is pierced by tiny foramina (Fig. 7C).</p><p>COMPARISONS</p><p>The holotype skull of Diplocynodon kochi n. sp. belongs to a mature individual as indicated by the presence of strongly sutured cranial bones and the deeply ornamented dorsal surfaces of the rostrum and the skull table.</p><p>The premaxillary surface in Diplocynodon kochi n. sp. possesses a deep notch lateral to naris, condition present in D. hantoniensis Wood, 1846, D. tormis Buscalioni et al., 1992 and Alligator Cuvier, 1807 (Rio et al. 2020), but also in D. ratelii (Díaz Aráez et al. 2017: fig. 6) and Orientalosuchus naduongensis Massonne et al., 2019 (Massonne et al. 2019). In the specimen of D. cf. hantoniensis, known from the late Eocene of Mormont, W-Switzerland (Pictet 1857), regarded as Alligatoroidea indet. by Rio et al. (2020), the rim of the premaxilla is extremely sharp and the external naris is comparatively large, features also comparable to D. kochi . The posterior premaxillary process of D. kochi is of the same length (i.e., reaching the level of third maxillary alveolus), as that of D. hantoniensis, D. muelleri Kälin, 1936 and D. ratelii, whereas that of D. remensis reaches only the level of the second maxillary alveolus (Martin et al. 2014). In contrast, in some Asian alligatoroids (e.g. Orientalosuchus naduongensis Massonne et al., 2019) the posterior premaxillary process extends posterior to the level of the fourth maxillary alveolus (Massone et al. 2019), whereas in some North American taxa (e.g. Brachychampsa), it may extend into the level of the fifth maxillary alveolus (Mook 1925). In the premaxilla of D. kochi the third and fourth alveoli are the largest, similar to D. hantoniensis and D. remensis (Martin et al. 2014) . Three occlusal pits are observed in the premaxillae of D. kochi positioned lingually to the alveoli, condition somewhat similar to D. hantoniensis (Rio et al. 2020), D. remensis (Martin et al. 2014) and D. muelleri (Piras &amp; Buscalioni 2006), whereas in D. tormis and D. ratelii the occlusal pits are interlocked between the premaxillary teeth (Piras &amp; Buscalioni 2006). The incisive foramen in D. kochi is larger than half of the greatest width of the premaxillae, whereas it is small in the remaining members of Diplocynodon . The nasals in D. kochi, similarly to most members of the genus Diplocynodon, are excluded from the naris, as it is seen also in Borealosuchus (Brochu 1999, 2000; Wu et al. 2001). In D. ratelii the nasals reach the external naris (Díaz Aráez et al. 2017; Luján et al. 2019), whereas in D. darwini the nasals almost contact the naris (Brochu 1999: fig. 32; Martin et al. 2014).</p><p>In D. kochi, the anteroposteriorly wide and mediolaterally shallow premaxillary-maxillary notch is bordered laterally by a bony ridge extending on the dorsolateral surface of the posterior premaxillary process. The resulting anterior margin of this notch stands roughly in an angle of 30° with the sagittal plane as opposed to the condition seen in some members of Diplocynodon (e.g. D. remensis) where the lateral margins of the premaxillae become perpendicular to the sagittal plane (Martin et al. 2014). The premaxillary-maxillary notch in D. muelleri, D. deponiae Frey et al., 1987 and D. hantoniensis is rather small or absent (Piras &amp; Buscalioni 2006; Delfino &amp; Smith 2012; Macaluso et al. 2019; Rio et al. 2020). In the remaining members of the genus, including D. darwini, D. ratelii, D. tormis, D. ungeri Prangner, 1845 and D. remensis the above constriction is present, but its morphology may be subject to ontogenetic variation. The bony ridge bordering the premaxillary-maxillary notch in D. kochi is positioned on the dorsolateral side of the premaxillary-maxillary notch, whereas in D. remensis two parallel ridges are positioned deeply within that notch, just posterior to the fifth premaxillary alveolus (Martin et al. 2014), and that part seemingly is remodelled ontogenetically by the enlarged dentary teeth.</p><p>The occlusion pits in the maxilla are distributed variably in the members of the genus, D. kochi resembling D. ratelii, D. tormis and D. muelleri, where the dentary teeth occlude in line with the maxillary toothrow (Díaz Aráez et al. 2017; Massone et al. 2019). The maxillary teeth and alveoli in D. kochi, up to the 11th tooth position, are more or less circular in shape, whereas the last five teeth and/or alveoli are slightly compressed mediolaterally, condition present also in D. remensis (Martin et al. 2014) and Brachychampsa montana Gilmore, 1911 (Mook 1925).</p><p>The anterior extent of the anterior process of the frontal in D. darwini is shorter than that of the jugal (Hastings &amp; Hellmund 2015: fig. 2), whereas it is approximately the same in D. hantoniensis, (e.g. in NHMUK OR 25170) (Rio et al. 2020), D. kochi (Fig. 3B) and D. tormis (Massone et al. 2019) . Nevertheless, the anterior extent of the jugal is shorter than that of the frontal in D. remensis (Martin et al. 2014: fig. 2), D. ratelii (Díaz Aráez et al. 2017: fig. 3) and D. muelleri (Massone et al. 2019) . The frontal of D. kochi lacks a preorbital step; it is also absent or weakly developed in D. hantoniensis, D. darwini, D. deponiae and D. ratelii (Rio et al. 2020), but it has been described in D. remensis (Martin et al. 2014), and figured in D. tormis (Buscalioni et al. 1992: fig. 2); the specimen STUS-344 of D. tormis apparently lacks any trace of this step in the preorbital area (Serrano-Martínez et al. 2019: fig. 1E).</p><p>The dorsal sculpture of D. kochi in the preorbital area approaches the condition seen in D. tormis, the latter possessing three short grooves: one is sagittal extending on the frontal process, while the other two extend anterolaterally, parallel with the orbital margins (Serrano-Martínez et al. 2019). However, the frontal process in the holotype of D. tormis, as reported by Buscalioni et al. (1992), is devoid of dorsal sculpture, probably representing an intraspecific variation. The preorbital ridge in D. kochi is wide but prominent and possesses a lateral overhang, however it is relatively short and restricted only to the surface of the lacrimal. In D. remensis a preorbital ridge has been reported by Martin et al. (2014: 877), but considered as weakly developed and therefore displaying the plesiomorphic condition. In Orientalosuchus naduongensis, the preorbital ridge is prominent, as it is in Krabisuchus Martin &amp; Lauprasert, 2010, Mourasuchus Price, 1964 and some crocodylids, and connected anteriorly to a prominent ridge developed on the maxilla (Massone et al. 2019).</p><p>The dorsolateral part of the postorbital bar is sculptured in D. kochi and D. hantoniensis, whereas in D. tormis that surface is entirely smooth and delimited dorsally by a sharp ridge (Buscalioni et al. 1992; Serrano-Martínez et al. 2019: fig. 1E).</p><p>The supratemporal margin of the parietal in D. kochi is produced into a wide overhang and the medial wall of the parietal is deeply inset forming a deep recess, probably serving as an extended shelf-like insertion surface for the specific jaw adductor muscles (MAMEP) being in control for rapid closure of the lower jaws in a palinal type jaw movement (Holliday &amp; Witmer 2007; Ősi 2014).</p><p>The morphology of the most posterior margin of the right maxilla and the imprint left by the left ectopterygoid on the medial side of left posterior maxillary ramus suggest that the ectopterygoid may have bordered only the last maxillary alveolus, whereas the next two alveoli were delimited by the maxilla, condition very similar to D. ratelii (Rio et al. 2020: fig. 29D).</p><p>The anterior process of the quadratojugal is short, extending as a triangular process onto the medial surface of the jugal, a character contrasting with the remaining members of Diplocynodon, where it is known (e.g. D. darwini, D. deponiae, D. ratelii, D. tormis). In D. kochi, the quadratojugal spine is not preserved, but the available part of the quadratojugal process extending along the posterior margin of the infratemporal fenestra remains more or less parallel suggesting that it has been devoid of this structure, as in other members of the genus.</p><p>The posterior margin of the otic aperture in D. kochi is inset similarly to D. darwini (Massone et al. 2019), D. hantoniensis (Rio et al. 2020: fig. 3A, B) and D. tormis (Serrano-Martínez et al. 2019: fig. 1A). This condition contrasts with that of the remaining members of the genus (if preserved), where the posterior margin of the otic aperture is flush with the paroccipital process, as it is observed in D. remensis (Martin et al. 2014: fig. 3A, B, D) and D. deponiae (Delfino &amp; Smith 2012: fig. 4).</p><p>The position of the lateral carotid foramen, situated above the dorsal margin of the basisphenoid, is similar to the condition known in D. tormis (Serrano-Martínez et al. 2019: figs 1D; 2F), but also to Asiatosuchus depressifrons Blainville, 1855 (Delfino &amp; Smith 2009) and crocodylids (see below). However, in the remaining members of the genus Diplocynodon (if known), the lateral carotid foramen opens laterally to the basisphenoid (Martin et al. 2014; Massone et al. 2019; Rio et al. 2020: fig. 3D), this trait representing the plesiomorphic condition.</p><p>PHYLOGENETIC ANALYSIS</p><p>Character-taxon matrix and analytical protocol</p><p>We added all the scored osteological characters of Diplocynodon kochi n. sp. (106 characters, representing 53% from the character list) to the character-taxon matrix (CTM) of Massone et al. (2019) consisting of 115 operational taxonomic units (OTU) and 202 phenotypic characters. The dataset of these authors, based on the original dataset of Brochu &amp; Storrs (2012), was amended and new characters were added from Wang et al. (2016), Cossette &amp; Brochu (2018) and Li et al. (2019), using Bernissartia fagesii Dollo, 1883 as the outgroup. The original CTM of Massone et al. (2019) was reduced by us to 77 OTUs (including D. kochi n. sp. and D. remensis) and to 199 phenotypic characters (i.e., without the last three characters, considered irrelevant for the present study). We also applied the emendations proposed to the original dataset by Massone et al. (2019: supplementary material); all the characters were treated as unordered. We performed a parsimony analysis using the TNT version 1.1 of Goloboff et al. (2008), in which the CTM was first analysed using the ‘New Tehnology search’ option with the sectorial search, ratchet, tree drift and tree fusing options as default parameters. A search for suboptimal trees ten steps longer than that of the most parsimonious tree was also completed to calculate the decay indices of Bremer (1994) and the common synapomorphies for all trees were also mapped in the ‘Optimize’ menu.</p></div>	https://treatment.plazi.org/id/2B04CD60072CFFE3FEA2C8D3F364F8BD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Venczel, Márton;Codrea, Vlad A.	Venczel, Márton, Codrea, Vlad A. (2022): A new late Eocene alligatoroid crocodyliform from Transylvania. Comptes Rendus Palevol 21 (20): 411-429, DOI: 10.5852/cr-palevol2022v21a20, URL: https://doi.org/10.5852/cr-palevol2022v21a20
