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BACKGROUND: Unenlagiine paravians are among the most relevant Gondwanan theropod dinosaur clades for understanding the origin of birds, yet their fossil record remains incomplete, with most taxa being represented by fragmentary material and/or separated by lengthy temporal gaps, frustrating attempts to characterize unenlagiine evolution. RESULTS AND CONCLUSIONS: Here we describe Diuqin lechiguanae gen. et sp. nov., a new unenlagiine taxon from the Upper Cretaceous (Santonian) Bajo de la Carpa Formation of the Neuquén Basin of Neuquén Province in northern Patagonia, Argentina that fills a substantial stratigraphic gap in the fossil record of these theropods. Although known only from a very incomplete postcranial skeleton, the preserved bones of Diuqin differ from corresponding elements in other unenlagiines, justifying the erection of the new taxon. Moreover, in several morphological aspects, the humerus of Diuqin appears intermediate between those of geologically older unenlagiines from the Neuquén Basin (e.g., Unenlagia spp. from the Turonian-Coniacian Portezuelo Formation) and that of the stratigraphically younger, larger-bodied Austroraptor cabazai from the Campanian-Maastrichtian Allen Formation. Consequently, the morphology of the new taxon appears to indicate a transitional stage in unenlagiine evolution. Phylogenetic analysis recovers Diuqin as a paravian with multiple plausible systematic positions, but the strongest affinity is with Unenlagiinae. The humerus of the new form exhibits subcircular punctures near its distal end that are interpreted as feeding traces most likely left by a conical-toothed crocodyliform, mammal, or theropod, the latter potentially corresponding to a megaraptorid or another unenlagiine individual. Thus, in addition to filling important morphological and temporal gaps in unenlagiine evolutionary history, the new taxon also offers information relating to the paleoecology of these theropods.

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Tetrapod limbs morphology is a reliable proxy of locomotor capacities. Beyond this, other aspects of life habits, such as predation abilities, can also be relevant to determine main morphofunctional appendicular properties, which ultimately reflect a compromise between different factors of the biological role. Dromaeosauridae is a dinosaur clade belonging to Theropoda, a group of bipedal predators. Dromaeosaurids represent an interesting study case, in which the hindlimbs have been proposed to be involved in both locomotion and predation activity. A peculiar feature characterizing all dromaeosaurids is a modified second pedal digit, which is typically related to predation. This theropod group is closely related to birds and diversified during the Cretaceous Period, mainly in the Northern Hemisphere (Laurasia). However, a subclade of dromaeosaurids, the Unenlagiinae, was recently recognized for Gondwana. Nevertheless, there are morphological differences between derived Laurasian dromaeosaurids (eudromaeosaurs) and unenlagiines. Such differences are observed in the proportions between hindlimb bones and in the presence of a subarctometatarsalian condition in unenlagiines, which is mainly characterized by a proximally constricted metatarsal III. To evaluate the function of these divergent morphologies, we conducted morphometric analyses and comparisons of qualitative morphological aspects, encompassing unenlagiines, other dromaeosaurids, as well as taxa from other theropod groups, including extant birds. The former approach consisted of two phylogenetic principal component analyses, one based on the main measurements of the hindlimb, and the other focused on the lengths of the pedal phalanges. The first analysis drew the unenlagiines close to taxa with long tibiae, as well as long and slender metatarsi. Instead, eudromaeosaurs are closer to taxa with shorter tibiae, and shorter and wider metatarsi. The second analysis showed that eudromaeosaurs and unenlagiines have similar phalangeal proportions, including the elongation of distal phalanges. However, the shorter second phalanx of the pedal digit II of eudromaeosaurs could have increased the force generated by this digit, which was the main predatory tool of the autopodium. This, together with a shorter and wider metatarsus, and a marked hinge-like morphology of the articular surfaces of metatarsals and phalanges, possibly allowed eudromaeosaurs to exert a great gripping strength and hunt large prey. Conversely, the longer and slender subarctometatarsus, and less well-marked hinge joints of unenlagiines possibly gave them greater cursorial capacities. Additionally, the longer second phalanx of digit II allowed unenlagiines fast movements of this digit to hunt smaller and elusive prey. Thus, the distinctive morphological evolutionary pathways of these two dromaeosaurid clades seem to have been influenced by the particular locomotor and predatory specializations that characterized each of these lineages.

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Here we provide a detailed description of the postcranial skeleton of the holotype and referred specimens of Buitreraptor gonzalezorum. This taxon was recovered as an unenlagiine dromaeosaurid in several recent phylogenetic studies and is the best represented Gondwanan dromaeosaurid discovered to date. It was preliminarily described in a brief article, but a detailed account of its osteology is emerging in recent works. The holotype is the most complete specimen yet found, so an exhaustive description of it provides much valuable anatomical information. The holotype and referred specimens preserve the axial skeleton, pectoral and pelvic girdles, and both fore- and hindlimbs. Diagnostic postcranial characters of this taxon include: anterior cervical centra exceeding the posterior limit of neural arch; eighth and ninth cervical vertebral centra with lateroventral tubercles; pneumatic foramina only in anteriormost dorsals; middle and posterior caudal centra with a complex of shallow ridges on lateral surfaces; pneumatic furcula with two pneumatic foramina on the ventral surface; scapular blade transversely expanded at mid-length; well-projected flexor process on distal end of the humerus; dorsal rim of the ilium laterally everted; and concave dorsal rim of the postacetabular iliac blade. A paleohistological study of limb bones shows that the holotype represents an earlier ontogenetic stage than one of the referred specimens (MPCA 238), which correlates with the fusion of the last sacral vertebra to the rest of the sacrum in MPCA 238. A revised phylogenetic analysis recovered Buitreraptor as an unenlagiine dromaeosaurid, in agreement with previous works. The phylogenetic implications of the unenlagiine synapomorphies and other characters, such as the specialized pedal digit II and the distal ginglymus on metatarsal II, are discussed within the evolutionary framework of Paraves.

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Over the past two decades, the record of South American unenlagiine dromaeosaurids was substantially increased both in quantity as well as in quality of specimens. Here is presented a summary review of the South American record for these theropods.Unenlagia comahuensis, Unenlagia paynemili, and Neuquenraptor argentinus come from the Portezuelo Formation, the former genus being the most complete and with putative avian features. Neuquenraptor is more incomplete and exhibits pedal features resembling those of Unenlagia. The earliest and most complete South American dromaeosaurid is Buitreraptor gonzalezoru, whose preserved cranial remains, provides important data in the characterization of unenlagiines. The most recently described,Austroraptor cabazai also with cranial remains, allows further comparisons with Laurasian lineages and a better characterization of unenlagiines. The possible synonymy between nenlagia and Neuquenraptor is discussed. Additional evidences from Brazil and Colombia show that dinosaurs with similar dentition to that of unenlagiines were present in the whole South America. However, it is not possible to discard that these remains may belong to other unknown maniraptoran lineages, considering the increasing number of taxa of this group found in South America.

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Over the past two decades, the record of South American unenlagiine dromaeosaurids was substantially increased both in quantity as well as in quality of specimens. Here is presented a summary review of the South American record for these theropods.Unenlagia comahuensis, Unenlagia paynemili, and Neuquenraptor argentinus come from the Portezuelo Formation, the former genus being the most complete and with putative avian features. Neuquenraptor is more incomplete and exhibits pedal features resembling those of Unenlagia. The earliest and most complete South American dromaeosaurid is Buitreraptor gonzalezoru, whose preserved cranial remains, provides important data in the characterization of unenlagiines. The most recently described,Austroraptor cabazai also with cranial remains, allows further comparisons with Laurasian lineages and a better characterization of unenlagiines. The possible synonymy between nenlagia and Neuquenraptor is discussed. Additional evidences from Brazil and Colombia show that dinosaurs with similar dentition to that of unenlagiines were present in the whole South America. However, it is not possible to discart that these remains may belong to other unknown maniraptoran lineages, considering the increasing number of taxa of this group found in South America.

Nas últimas duas décadas, o registro do dromaeossaurídeos Unenlagiinae da América do Sul aumentou substancialmente tanto em quantidade, assim como na qualidade dos espécimes. Uma revisão sumária do registro sul americano para estes terópodos é apresentada aqui. Unenlagia comahuensis, Unenlagia paynemili e Neuquenraptor argentinus provenientes da Formação Portezuelo, sendo o primeiro o mais completo e com características avianas putativas. Neuquenraptor, muito incompleto, demonstra as características do pé que assemelhamse àquelas de Unenlagia. O mais velho, e mais completo dromaeossaurídeo americano é Buitreraptor gonzalezorum, cujo possui preservado ossos cranianos, fornece implicações importantes na caracterização dos unenlagiines. Descrito recentemente, Austroraptor cabazai também com ossos cranianos, permite mais comparações com linhagens provenientes da Laurasia e uma melhor caracterização dos unenlagiineos. A possível sinonímia entre Unenlagia e Neuquenraptor é discutido. As evidências adicionais do Brasil e da Colômbia mostram que dinossauros com dentição semlhante à dos unenlagiineos eram representados em toda América do Sul; entretanto, estas evidências podem igualmente ser relacionadas a outros maniraptores desconhecidos, considerando a grande diversidade do grupo na América do Sul.

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