An extinct deep-snouted Alligator species from the Quaternary of Thailand and comments on the evolution of crushing dentition in alligatorids.

Fossil Alligator remains from Asia are critical for tracing the enigmatic evolutionary origin of the Chinese alligator, Alligator sinensis, the only living representative of Alligatoridae outside the New World. The Asian fossil record is extremely scarce and it remains unknown whether A. sinensis is an anagenetic lineage or alternatively, extinct divergent species were once present. We provide a detailed comparative description of a morphologically highly distinct Alligator skull from the Quaternary of Thailand. Several autapomorphic characters warrant the designation of a new species. Alligator munensis sp. nov. shares obvious derived features with A. sinensis but autapomorphies imply a cladogenetic split, possibly driven by the uplift of the southeastern Tibetan plateau. The presence of enlarged posterior alveoli in Alligator munensis is most consistent with a reversal to the alligatorine ancestral condition of having crushing dentition, a morphology strikingly absent among living alligatorids. Crushing dentition has been previously considered to indicate an ecological specialisation in early alligatorines that was subsequently lost in Alligator spp. However, we argue that there is yet no evidence for crushing dentition reflecting an adaptation for a narrower niche, while opportunistic feeding, including seasonal utilisation of hard-shelled preys, is a reasonable alternative interpretation of its function.

A new specimen of Solnhofia parsonsi from the Upper Jurassic (Kimmeridgian) Plattenkalk deposits of Painten (Bavaria, Germany) and comments on the relationship between limb taphonomy and habitat ecology in fossil turtles.

The limestones of the Solnhofen area in southern Germany are one of the most important fossil Lagerstätten from the entire Mesozoic era, especially famous for the exquisitely preserved vertebrates. The turtles from the Solnhofen Limestone have been always of special interest because they include some of the best-preserved specimens from the Mesozoic. Here, we describe a new turtle specimen from the Torleite Formation (Kimmeridgian) of Painten and refer it to the thalassochelydian turtle Solnhofia parsonsi based on the presence of a unique combination of characters. The far majority of morphological differences from previously published specimens can be explained by ontogeny as the new specimen represents a larger, more ossified, and presumably older individual. Additionally, the specimen from Painten is the first described specimen of S. parsonsi preserving the largely complete and articulated limbs, the preservation of which indicates that the taxon did not possess stiffened paddles present in more pelagic marine turtles and is consistent with a previously inferred nearshore marine lifestyle. Contrary to previous inferences, we argue that taphonomic preservation of digits in articulated fossil turtles from laminated deposits cannot be used alone to infer marine or freshwater habitat. Finally, the new specimen from Painten is only the second, for which detailed information on its stratigraphic position and locality of origin are known.

A forged 'chimera' including the second specimen of the protostegid sea turtle Santanachelysgaffneyi and shell parts of the pleurodire Araripemys from the Lower Cretaceous Santana Group of Brazil.

Fossils of Cretaceous sea turtles adapted to an open marine lifestyle remain rare finds to date. Furthermore, the relationships between extant sea turtles, chelonioids, and other Mesozoic marine turtles are still contested, with one key species being Santanachelys gaffneyi Hirayama, 1998, long considered the earliest true sea turtle. The species is an Early Cretaceous member of Protostegidae, a controversial clade either placed within or closely related to Chelonioidea or, alternatively, along the stem lineage of hidden-neck turtles (Cryptodira) and representing an independent open marine radiation. Santanachelys gaffneyi is one of the most completely preserved early protostegids and is therefore critical for establishing the global phylogenetic position of the group. However, the single known specimen of this taxon is yet to be described in detail. Here we describe a second specimen of Santanachelys gaffneyi from its type horizon, the Romualdo Formation (late Aptian) of the Santana Group of the Araripe basin, NE Brazil. The skeletal elements preserved include the posterior part of the skull, neck vertebrae, shoulder girdle, anterior-most and left/central part of the carapace with few peripherals, and plastron lacking most of the hyoplastra. The remaining part of the carapace was apparently completed by fossil dealers using an anterior part of the pleurodiran Araripemydidae, tentatively identified as a shell portion of cf. Araripemys barretoi, a more common Santana fossil turtle, among other indeterminate turtle shell fragments. The purpose of this paper is to report the repatriation of the specimen to Brazil and to provide a preliminary description. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-023-00271-9.

The impact of molecular data on the phylogenetic position of the putative oldest crown crocodilian and the age of the clade.

The use of molecular data for living groups is vital for interpreting fossils, especially when morphology-only analyses retrieve problematic phylogenies for living forms. These topological discrepancies impact on the inferred phylogenetic position of many fossil taxa. In Crocodylia, morphology-based phylogenetic inferences differ fundamentally in placing Gavialis basal to all other living forms, whereas molecular data consistently unite it with crocodylids. The Cenomanian Portugalosuchus azenhae was recently described as the oldest crown crocodilian, with affinities to Gavialis, based on morphology-only analyses, thus representing a potentially important new molecular clock calibration. Here, we performed analyses incorporating DNA data into these morphological datasets, using scaffold and supermatrix (total evidence) approaches, in order to evaluate the position of basal crocodylians, including Portugalosuchus. Our analyses incorporating DNA data robustly recovered Portugalosuchus outside Crocodylia (as well as thoracosaurs, planocraniids and Borealosuchus spp.), questioning the status of Portugalosuchus as crown crocodilian and any future use as a node calibration in molecular clock studies. Finally, we discuss the impact of ambiguous fossil calibration and how, with the increasing size of phylogenomic datasets, the molecular scaffold might be an efficient (though imperfect) approximation of more rigorous but demanding supermatrix analyses.

Divergent mammalian body size in a stable Eocene greenhouse climate.

A negative correlation between body size and the latitudinal temperature gradient is well established for extant terrestrial endotherms but less so in the fossil record. Here we analyze the middle Eocene site of Geiseltal (Germany), whose record is considered to span ca. 5 Myrs of gradual global cooling, and generate one of the most extensive mammalian Paleogene body size datasets outside North America. The δ18O and δ13C isotopic analysis of bioapatite reveals signatures indicative of a humid, subtropical forest with no apparent climatic change across Geiseltal. Yet, body mass of hippomorphs and tapiromorphs diverges rapidly from a respective median body size of 39 kg and 124 kg at the base of the succession to 26 kg and 223 kg at the top. We attribute the divergent body mass evolution to a disparity in lifestyle, in which both taxa maximize their body size-related selective advantages. Our results therefore support the view that intrinsic biotic processes are an important driver of body mass outside of abrupt climate events. Moreover, the taxonomy previously used to infer the duration of the Geiseltal biota is not reproducible, which precludes chronological correlation with Eocene marine temperature curves.

A new alligatoroid from the Eocene of Vietnam highlights an extinct Asian clade independent from extant Alligator sinensis.

During systematic paleontological surveys in the Na Duong Basin in North Vietnam between 2009 and 2012, well-preserved fossilized cranial and postcranial remains belonging to at least 29 individuals of a middle to late Eocene (late Bartonian to Priabonian age (39-35 Ma)) alligatoroid were collected. Comparative anatomical study of the material warrants the diagnosis of a new taxon, Orientalosuchus naduongensis gen. et sp. nov. The combined presence of an enlarged fifth maxillary tooth, prominent preorbital ridges, a large supraoccipital exposure on the skull table, a palatine-pterygoid suture anterior to the posterior end of the suborbital fenestra, and a pterygoid forming a neck surrounding the choana is unique to this species. Unlike previous phylogenies, our parsimony analysis recovers a monophyletic Late Cretaceous to Paleogene East to Southeastern Asian alligatoroid group, here named Orientalosuchina. The group includes Orientalosuchus naduongensis, Krabisuchus siamogallicus, Eoalligator chunyii, Jiangxisuchus nankangensis and Protoalligator huiningensis, all of them sharing a medial shifted quadrate foramen aerum. The recognition of this clade indicates at least two separate dispersal events from North America to Asia: one during the Late Cretaceous by Orientalosuchina and one by the ancestor of Alligator sinensis during the Paleogene or Neogene, the timing of which is poorly constrained.

The sinemydid turtle Ordosemys from the Lower Cretaceous Mengyin Formation of Shandong, China and its implication for the age of the Luohandong Formation of the Ordos Basin.

Chronostratigraphic correlation of terrestrial Early Cretaceous biotas in China is highly problematic due to the lack of marine deposits, few absolute dates, and limited number of index fossils. This often leaves vertebrate faunas as one of the few potential tools for a preliminary biostratigraphy. Taxonomic identity of fragmentary fossils is, however, often uncertain and many faunas are insufficiently sampled. Turtles are one of the most common elements of Early Cretaceous biotas of Asia and their skeleton is frequently preserved more completely than that of other vertebrates- they yet receive little attention from vertebrate paleontologists. We here record the presence of the sinemydid turtle Ordosemys leios from the Lower Cretaceous Mengyin Formation of Shandong Province, China, best known for the first dinosaurs and Mesozoic turtles described from the country. Ordosemys is the third turtle reported from the Mengyin Formation along with Sinemys lens and Sinochelys applanata and the only other formation where Ordosemys is known to co-occur with Sinemys is the Luohandong Formation of the Ordos Basin (Inner Mongolia), the type and so far only horizon of Ordosemys leios. The presence of the crocodyliform Shantungosuchus may further define a fauna that is so far only known from these two formations. The stratigraphic position of the Luohandong Formation is poorly controlled and it has been placed anywhere between the Valanginian and Aptian. Published absolute dates from the Mengyin Formation and the numerous shared vertebrate and invertebrate taxa (now also including turtles) implies a Valanginian-early Hauterivian age for the Luohandong Formation-in contrast to late Hauterivian-Albian as previously proposed using the temporal distribution of Psittacosaurus. The new specimen of Ordosemys leios preserves the only known manus of this species and ecomorphological analysis of limb proportions implies that it was a less capable swimmer compared to Ordosemys liaoxiensis coming from the younger Jehol Biota.

A new large-bodied thalattosuchian crocodyliform from the Lower Jurassic (Toarcian) of Hungary, with further evidence of the mosaic acquisition of marine adaptations in Metriorhynchoidea.

Based on associated and three-dimensionally preserved cranial and postcranial remains, a new thalattosuchian crocodyliform, Magyarosuchus fitosi gen. et sp. nov. from the Lower Jurassic (Upper Toarcian) Kisgerecse Marl Formation, Gerecse Mountains, Hungary is described here. Phylogenetic analyses using three different datasets indicate that M. fitosi is the sister taxon of Pelagosaurus typus forming together the basal-most sub-clade of Metriorhynchoidea. With an estimated body length of 4.67-4.83 m M. fitosi is the largest known non-metriorhynchid metriorhynchoid. Besides expanding Early Jurassic thalattosuchian diversity, the new specimen is of great importance since, unlike most contemporaneous estuarine, lagoonal or coastal thalattosuchians, it comes from an 'ammonitico rosso' type pelagic deposit of the Mediterranean region of the Tethys. A distal caudal vertebra having an unusually elongate and dorsally projected neural spine implies the presence of at least a rudimentary hypocercal tail fin and a slight ventral displacement of the distal caudal vertebral column in this basal metriorhynchoid. The combination of retaining heavy dorsal and ventral armors and having a slight hypocercal tail is unique, further highlighting the mosaic manner of marine adaptations in Metriorhynchoidea.

Total evidence analysis and body size evolution of extant and extinct tortoises (Testudines: Cryptodira: Pan-Testudinidae).

Testudinidae (tortoises) is an extant clade of terrestrial turtles of worldwide distribution and with a rich fossil record that provides an exceptional context for studying their evolutionary history. Because of the lack of global phylogenetic analyses integrating extinct taxa, our current knowledge of the relationships of the total clade of Testudinidae is rather poor. To resolve this issue, we performed the first total evidence analysis of Pan-Testudinidae. The total evidence trees are congruent with the molecular topology and agree on the dichotomy of derived Testudinidae (=Testudininae; Converted Clade Name) into two previously recognized major clades, Testudona and Geochelona (New Clade Name). The integration of extinct taxa into the analysis allowed the stratigraphic fit of the total evidence trees, indicating that crown Testudininae, Testudona and Geochelona all originated by the Late Eocene, in agreement with recent molecular estimates. Ghost lineage analysis indicates high diversification in the Late Eocene and in the Miocene. The age of crown Testudo is Late Miocene, again in accordance with some molecular dates. Phylogenetic placement of fossils demonstrates that giant body size independently evolved in multiple continental mainland taxa and confirms recent results deduced from living taxa-giantism in Testudinidae is not linked to the insular effect. An unexpected outcome is the recovery of miniaturization in Testudona (<30 cm carapace length) that emerged sometime between the Oligocene and Early Miocene. No clear correlation between body size evolution and climate is apparent, but increased taxon sampling may nevertheless demonstrate the role of cooling and warming as one of many influential variables.

Lower Cretaceous fossils from China shed light on the ancestral body plan of crown softshell turtles (Trionychidae, Cryptodira).

Pan-trionychids or softshell turtles are a highly specialized and widespread extant group of aquatic taxa with an evolutionary history that goes back to the Early Cretaceous. The earliest pan-trionychids had already fully developed the "classic" softshell turtle morphology and it has been impossible to resolve whether they are stem members of the family or are within the crown. This has hindered our understanding of the evolution of the two basic body plans of crown-trionychids. Thus it remains unclear whether the more heavily ossified shell of the cyclanorbines or the highly reduced trionychine morphotype is the ancestral condition for softshell turtles. A new pan-trionychid from the Early Cretaceous of Zhejiang, China, Perochelys hengshanensis sp. nov., allows a revision of softshell-turtle phylogeny. Equal character weighting resulted in a topology that is fundamentally inconsistent with molecular divergence date estimates of deeply nested extant species. In contrast, implied weighting retrieved Lower Cretaceous Perochelys spp. and Petrochelys kyrgyzensis as stem trionychids, which is fully consistent with their basal stratigraphic occurrence and an Aptian-Santonian molecular age estimate for crown-trionychids. These results indicate that the primitive morphology for soft-shell turtles is a poorly ossified shell like that of crown-trionychines and that shell re-ossification in cyclanorbines (including re-acquisition of peripheral elements) is secondary.

A toothed turtle from the Late Jurassic of China and the global biogeographic history of turtles.

BACKGROUND: Turtles (Testudinata) are a successful lineage of vertebrates with about 350 extant species that inhabit all major oceans and landmasses with tropical to temperate climates. The rich fossil record of turtles documents the adaptation of various sub-lineages to a broad range of habitat preferences, but a synthetic biogeographic model is still lacking for the group. RESULTS: We herein describe a new species of fossil turtle from the Late Jurassic of Xinjiang, China, Sichuanchelys palatodentata sp. nov., that is highly unusual by plesiomorphically exhibiting palatal teeth. Phylogenetic analysis places the Late Jurassic Sichuanchelys palatodentata in a clade with the Late Cretaceous Mongolochelys efremovi outside crown group Testudines thereby establishing the prolonged presence of a previously unrecognized clade of turtles in Asia, herein named Sichuanchelyidae. In contrast to previous hypotheses, M. efremovi and Kallokibotion bajazidi are not found within Meiolaniformes, a clade that is here reinterpreted as being restricted to Gondwana. CONCLUSIONS: A revision of the global distribution of fossil and recent turtle reveals that the three primary lineages of derived, aquatic turtles, including the crown, Paracryptodira, Pan-Pleurodira, and Pan-Cryptodira can be traced back to the Middle Jurassic of Euramerica, Gondwana, and Asia, respectively, which resulted from the primary break up of Pangaea at that time. The two primary lineages of Pleurodira, Pan-Pelomedusoides and Pan-Chelidae, can similarly be traced back to the Cretaceous of northern and southern Gondwana, respectively, which were separated from one another by a large desert zone during that time. The primary divergence of crown turtles was therefore driven by vicariance to the primary freshwater aquatic habitat of these lineages. The temporally persistent lineages of basal turtles, Helochelydridae, Meiolaniformes, Sichuanchelyidae, can similarly be traced back to the Late Mesozoic of Euramerica, southern Gondwana, and Asia. Given the ambiguous phylogenetic relationships of these three lineages, it is unclear if their diversification was driven by vicariance as well, or if they display a vicariance-like pattern. The clean, primary signal apparent among early turtles is secondarily obliterated throughout the Late Cretaceous to Recent by extensive dispersal of continental turtles and by multiple invasions of marine habitats.

A sinemydid turtle from the Jehol Biota provides insights into the basal divergence of crown turtles.

Morphological phylogenies stand in a major conflict with molecular hypotheses regarding the phylogeny of Cryptodira, the most diverse and widely distributed clade of extant turtles. However, molecular hypotheses are often considered a better estimate of phylogeny given that it is more consistent with the stratigraphic and geographic distribution of extinct taxa. That morphology fails to reproduce the molecular topology partly originates from problematic character polarization due to yet another contradiction around the composition of the cryptodiran stem lineage. Extinct sinemydids are one of these problematic clades: they have been either placed among stem-cryptodires, stem-chelonioid sea turtles, or even stem-turtles. A new sinemydid from the Early Cretaceous Jehol Biota (Yixian Formation, Barremian-Early Aptian) of China, Xiaochelys ningchengensis gen. et sp. nov., allows for a reassessment of the phylogenetic position of Sinemydidae. Our analysis indicates that sinemydids mostly share symplesiomorphies with sea turtles and their purported placement outside the crown-group of turtles is an artefact of previous datasets. The best current phylogenetic estimate is therefore that sinemydids are part of the stem lineage of Cryptodira together with an array of other Jurassic to Cretaceous taxa. Our study further emphasises the importance of using molecular scaffolds in global turtle analyses.

An enigmatic crocodyliform tooth from the bauxites of western Hungary suggests hidden mesoeucrocodylian diversity in the Early Cretaceous European archipelago.

Background. The Cretaceous of southern Europe was characterized by an archipelago setting with faunas of mixed composition of endemic, Laurasian and Gondwanan elements. However, little is known about the relative timing of these faunal influences. The Lower Cretaceous of East-Central Europe holds a great promise for understanding the biogeographic history of Cretaceous European biotas because of the former proximity of the area to Gondwana (as part of the Apulian microcontinent). However, East-Central European vertebrates are typically poorly known from this time period. Here, we report on a ziphodont crocodyliform tooth discovered in the Lower Cretaceous (Albian) Alsópere Bauxite Formation of Olaszfalu, western Hungary. Methods. The morphology of the tooth is described and compared with that of other similar Cretaceous crocodyliforms. Results. Based on the triangular, slightly distally curved, constricted and labiolingually flattened crown, the small, subequal-sized true serrations on the carinae mesially and distally, the longitudinal fluting labially, and the extended shelves along the carinae lingually the tooth is most similar to some peirosaurid, non-baurusuchian sebecosuchian, and uruguaysuchid notosuchians. In addition, the paralligatorid Wannchampsus also possesses similar anterior teeth, thus the Hungarian tooth is referred here to Mesoeucrocodylia indet. Discussion. Supposing a notosuchian affinity, this tooth is the earliest occurrence of the group in Europe and one of the earliest in Laurasia. In case of a paralligatorid relationship the Hungarian tooth would represent their first European record, further expanding their cosmopolitan distribution. In any case, the ziphodont tooth from the Albian bauxite deposit of western Hungary belongs to a group still unknown from the Early Cretaceous European archipelago and therefore implies a hidden diversity of crocodyliforms in the area.

A new specimen of Manchurochelys manchoukuoensis from the Early Cretaceous Jehol Biota of Chifeng, Inner Mongolia, China and the phylogeny of Cretaceous basal eucryptodiran turtles.

BACKGROUND: Manchurochelys manchoukuoensis is an emblematic turtle from the Cretaceous Yixian Formation of Liaoning, China, a geological rock unit that is famous for yielding perfectly preserved skeletons of fossil vertebrates, including that of feathered dinosaurs. Manchurochelys manchoukuoensis was one of the first vertebrates described from this fauna, also known as the Jehol Biota. The holotype was lost during World War II and only one additional specimen has been described since. Manchurochelys manchoukuoensis is a critical taxon for unraveling the phylogenetic relationships of Cretaceous pancryptodires from Asia, a group that is considered to be of key importance for the origin of crown-group hidden-neck turtles (Cryptodira). RESULTS: A new specimen of Manchurochelys manchoukuoensis is described here from the Jiufotang Formation of Qilinshan, Chifeng, Inner Mongolia, China. This is the third specimen described and expands the range of this taxon from the Yixian Formation of the Fuxin-Yixian Basin in Liaoning to the Jiufotang Formation of the Chifeng-Yuanbaoshan Basin. A possible temporal extension of the range is less certain. The new finding adds to our understanding of the morphology of this taxon and invites a thorough revision of the phylogeny of Macrobaenidae, Sinemydidae, and closely allied forms. CONCLUSIONS: Our comprehensive phylogenetic analyses of Cretaceous Asian pancryptodires yielded two main competing hypotheses: in the first these taxa form a paraphyletic grade, whereas in the second they form a monophyletic clade. The inclusion of problematic tree changing taxa, such as Panpleurodires (stem + crown side-neck turtles) has a major influence on the phylogenetic relationships of Sinemydidae and closely allied forms. Manchurochelys manchoukuoensis nests within Sinemydidae together with Sinemys spp. and Dracochelys bicuspis in the majority of our analyses.

A new xinjiangchelyid turtle from the Middle Jurassic of Xinjiang, China and the evolution of the basipterygoid process in Mesozoic turtles.

BACKGROUND: Most turtles from the Middle and Late Jurassic of Asia are referred to the newly defined clade Xinjiangchelyidae, a group of mostly shell-based, generalized, small to mid-sized aquatic froms that are widely considered to represent the stem lineage of Cryptodira. Xinjiangchelyids provide us with great insights into the plesiomorphic anatomy of crown-cryptodires, the most diverse group of living turtles, and they are particularly relevant for understanding the origin and early divergence of the primary clades of extant turtles. RESULTS: Exceptionally complete new xinjiangchelyid material from the ?Qigu Formation of the Turpan Basin (Xinjiang Autonomous Province, China) provides new insights into the anatomy of this group and is assigned to Xinjiangchelys wusu n. sp. A phylogenetic analysis places Xinjiangchelys wusu n. sp. in a monophyletic polytomy with other xinjiangchelyids, including Xinjiangchelys junggarensis, X. radiplicatoides, X. levensis and X. latiens. However, the analysis supports the unorthodox, though tentative placement of xinjiangchelyids and sinemydids outside of crown-group Testudines. A particularly interesting new observation is that the skull of this xinjiangchelyid retains such primitive features as a reduced interpterygoid vacuity and basipterygoid processes. CONCLUSIONS: The homology of basipterygoid processes is confidently demonstrated based on a comprehensive review of the basicranial anatomy of Mesozoic turtles and a new nomenclatural system is introduced for the carotid canal system of turtles. The loss of the basipterygoid process and the bony enclosure of the carotid circulation system occurred a number of times independently during turtle evolution suggesting that the reinforcement of the basicranial region was essential for developing a rigid skull, thus paralleling the evolution of other amniote groups with massive skulls.

An enormous Jurassic turtle bone bed from the Turpan Basin of Xinjiang, China.

A spectacular new terrestrial Konzentratlagerstätte is introduced from the Turpan Basin of Xinjiang, China that probably belongs to the late Middle Jurassic Qigu Formation. It contains a mass accumulation of "xinjiangchelyid" turtles preliminarily identified as Annemys sp. In the zone with the highest turtle concentration, complete and articulated turtle skeletons are tightly packed at a density of up to 36 turtles per square meter. The fossiliferous layer is thickened here and shows an erosional base. This high concentration zone outcrops approximately 10 m in length and shows no decrease in turtle density after exposing 2 m of the layer into the hillside. Adjacent is a more expansive zone of at least 10 m by 30 m. In this region, the fossiliferous layer is evenly thick, and approximately five, fully disarticulated turtles are present per square meter. A conservatively estimated 1,800 turtles may, therefore, have been deposited at this site. It is likely that these aquatic turtles gathered in a retreating water hole in a riverine environment during a drought, much as some aquatic turtles will do today, but perished when the habitat dried up completely. A following catastrophic rainfall event caused a debris flow, possibly channelized in a dry river bed, which transported complete turtles, disarticulated turtles, and mudstone clasts and deposited them after a short distance. This taphonomic model is consistent with previous environmental reconstructions of the Turpan Basin during the late Middle Jurassic in predicting the episodic breakdown of regional monsoonal circulation resulting in a seasonally dry climate with severe episodic droughts.

Survival of Theriosuchus (Mesoeucrocodylia: Atoposauridae) in a Late Cretaceous archipelago: a new species from the Maastrichtian of Romania.

Small terrestrial non-eusuchian mesoeucrocodylians are common components of Cretaceous assemblages of Gondwanan provinces with notosuchians and araripesuchids as flagship taxa in South America, Africa and Madagascar, well into the Late Cretaceous. On the other hand, these are exceedingly rare in Laurasian landmasses during the Late Cretaceous. Small terrestrial mesoeucrocodylians from Europe were often referred to the genus Theriosuchus, a taxon with stratigraphic range extending from the Late Jurassic to the late Early Cretaceous. Theriosuchus is abundantly reported from various European localities, although Asiatic and possibly North American members are also known. It has often been closely associated with the first modern crocodilians, members of the Eusuchia, because of the presence of procoelous vertebrae, a widespread key character diagnosing the Eusuchia. Nevertheless, the relationships of Theriosuchus have not been explored in detail although one species, Theriosuchus pusillus, has been extensively described and referred in numerous works. Here, we describe a new basal mesoeucrocodylian, Theriosuchus sympiestodon sp. nov. from the Maastrichtian of the Hateg Basin, Romania, suggesting a large temporal gap (about 58 myr) in the fossil record of the genus. Inclusion of the new taxon, along with Theriosuchus guimarotae, in a phylogenetic analysis confirms its referral to the genus Theriosuchus, within a monophyletic atoposaurid clade. Although phylogenetic resolution within this clade is still poor, the new taxon appears, on morphological grounds, to be most closely related to T. pusillus. The relationships of Atoposauridae within Mesoeucrocodylia and especially to Neosuchia are discussed in light of the results of the present contribution as well as from recent work. Our results raise the possibility that Atoposauridae might not be regarded as a derived neosuchian clade anymore, although further investigation of the neosuchian interrelationships is needed. Reports of isolated teeth referable to a closely related taxon from the Upper Cretaceous of Romania and France, together with the presence of Doratodon and Ischyrochampsa, indicate a previously unsuspected diverse assemblage of non-eusuchian mesoeucrocodylians in the Late Cretaceous European archipelago.