RESUMEN
While brain size in primates and their relatives within Euarchontoglires is well-studied, less research has examined brain shape, or the allometric trajectories that underlie the relationship between size and shape. Defining these patterns is key to understanding evolutionary trends. 3D geometric morphometric analyses of endocranial shape were performed on 140 species of extant euarchontoglirans using digital cranial endocasts. Principal component analyses on Procrustes shape variables show a clear phylogenetic pattern in endocranial shape, supported by an ANOVA which identified significant differences in shape among several groups (e.g., Platyrrhini, Strepsirrhini, Scandentia, Rodentia, and Lagomorpha). ANOVAs of shape and size also indicate that allometry has a small but significant impact on endocranial shape across Euarchontoglires, with homogeneity of slopes tests finding significant differences in the scaling relationship between shape and size among these same groups. While most of these clades possess a distinct endocranial morphotype, the highly derived platyrrhines display the strongest relationship between size and shape. Rodents show the most diversity in endocranial shape, potentially attributed to their comparatively weak relationship between shape and size. These results suggest fundamental differences in how shape and size covary among Euarchontoglires, which may have facilitated the adaptive radiations that characterize members of this group.
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Evolución Biológica , Filogenia , Cráneo , Animales , Cráneo/anatomía & histología , Fósiles/anatomía & histología , Análisis de Componente Principal , Encéfalo/anatomía & histología , Primates/anatomía & histologíaRESUMEN
Tardigrades are a diverse phylum of microscopic invertebrates widely known for their extreme survival capabilities. Molecular clocks suggest that tardigrades diverged from other panarthropods before the Cambrian, but their fossil record is extremely sparse. Only the fossil tardigrades Milnesium swolenskyi (Late Cretaceous) and Paradoryphoribius chronocaribbeus (Miocene) have resolved taxonomic positions, restricting the availability of calibration points for estimating for the origin of this phylum. Here, we revise two crown-group tardigrades from Canadian Cretaceous-aged amber using confocal fluorescence microscopy, revealing critical morphological characters that resolve their taxonomic positions. Formal morphological redescription of Beorn leggi reveals that it features Hypsibius-type claws. We also describe Aerobius dactylus gen. et sp. nov. based on its unique combination of claw characters. Phylogenetic analyses indicate that Beo. leggi and Aer. dactylus belong to the eutardigrade superfamily Hypsibioidea, adding a critical fossil calibration point to investigate tardigrade origins. Our molecular clock estimates suggest an early Paleozoic diversification of crown-group Tardigrada and highlight the importance of Beo. leggi as a calibration point that directly impacts estimates of shallow nodes. Our results suggest that independent terrestrialization of eutardigrades and heterotardigrades occurred around the end-Carboniferous and Lower Jurassic, respectively. These estimates also provide minimum ages for convergent acquisition of cryptobiosis.
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Ámbar , Evolución Biológica , Fósiles , Filogenia , Tardigrada , Animales , Fósiles/anatomía & histología , Tardigrada/clasificación , Tardigrada/anatomía & histología , Tardigrada/genética , CanadáRESUMEN
Fossil identification practices have a profound effect on our interpretation of the past because these identifications form the basis for downstream analyses. Therefore, well-supported fossil identifications are necessary for examining the impact of past environmental changes on populations and communities. Here we apply an apomorphic identification framework in a case study identifying fossil lizard remains from Hall's Cave, a late Quaternary fossil site located in Central Texas, USA. We present images and descriptions of a broad comparative sample of North American lizard cranial elements and compile new and previously reported apomorphic characters for identifying fossil lizards. Our fossil identifications from Hall's Cave resulted in a minimum of 11 lizard taxa, including five lizard taxa previously unknown from the site. Most of the identified fossil lizard taxa inhabit the area around Hall's Cave today, but we reinforce the presence of an extirpated species complex of horned lizard. A main goal of this work is to establish a procedure for making well-supported fossil lizard identifications across North America. The data from this study will assist researchers endeavoring to identify fossil lizards, increasing the potential for novel discoveries related to North American lizards and facilitating more holistic views of ancient faunal assemblages.
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Fósiles , Lagartos , Cráneo , Animales , Fósiles/anatomía & histología , Lagartos/anatomía & histología , Cráneo/anatomía & histología , Texas , CuevasRESUMEN
The separation of closely related terrestrial or freshwater species by vast marine barriers represents a biogeographical riddle. Such cases can provide evidence for vicariance, a process whereby ancient geological events like continental rifting divided ancestral geographical ranges. With an evolutionary history extending tens of millions of years, freshwater ecology, and distribution encompassing widely separated southern landmasses, osteoglossid bonytongue fishes are a textbook case of vicariance attributed to Mesozoic fragmentation of the Gondwanan supercontinent. Largely overlooked fossils complicate the clean narrative invoked for extant species by recording occurrences on additional continents and in marine settings. Here, we present a new total-evidence phylogenetic hypothesis for bonytongue fishes combined with quantitative models of range evolution and show that the last common ancestor of extant osteoglossids was likely marine, and that the group colonized freshwater settings at least four times when both extant and extinct lineages are considered. The correspondence between extant osteoglossid relationships and patterns of continental fragmentation therefore represents a striking example of biogeographical pseudocongruence. Contrary to arguments against vicariance hypotheses that rely only on temporal or phylogenetic evidence, these results provide direct palaeontological support for enhanced dispersal ability early in the history of a group with widely separated distributions in the modern day.
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Distribución Animal , Evolución Biológica , Peces , Fósiles , Filogenia , Animales , Fósiles/anatomía & histología , Peces/anatomía & histologíaRESUMEN
Constraint is a fundamental concept in evolutionary theory. Morphology and ecology both are limited by functional, historical and developmental factors to a subset of the theoretical range species could occupy. Cat-like carnivorans (Feliformia) offer a unique opportunity to investigate phenotypic constraint, as several feliform clades are purported to be limited to generalized ecomorphological roles, while others possess extremely specialized durophagous (bone-crushing) and sabretooth morphology. We investigated the evolutionary history of feliforms by considering their phylogeny, morphological disparity and rates of evolution. We recover results that show a mosaic pattern exists in the degree of morphological disparity per anatomical region per clade and ecology. Non-hypercarnivores, such as viverrids (civets and genets), Malagasy euplerids and lophocyonids (extinct hypocarnivores), have the greatest dental disparity, while hypercarnivores (felids, nimravids, many hyaenids) have the lowest dental disparity but highest cranial and mandibular disparity (excluding dentition). However, high disparity is not necessarily associated with high rates of evolution, but instead with ecological radiations. We reveal that relationships between specialization and disparity are not as simple as past research has concluded. Instead, morphological disparity results from an anatomical mosaic of evolution, where different ecologies correlate with and likely channel unique patterns/combinations of disparity per anatomical partition.
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Evolución Biológica , Filogenia , Animales , Cráneo/anatomía & histología , Fósiles/anatomía & histologíaRESUMEN
The record of Mesozoic reptiles in the Northern region of Brazil is extremely limited, with the only definite occurrence consisting of two ziphodont teeth recovered from an oil well core in the municipality of Nova Olinda do Norte, Amazonas state, from strata of the Alter do Chão Formation (Cenomanian) of the Amazonas Basin. In this study, we aim to reevaluate MCT.R.514 using the most recent methodologies available for the identification of isolated theropod teeth. The cladistic analyses recovered MCT.R.514 as a possible metriacanthosaurid or non carcharodontosaurid allosauroid while the LDAs showed affinities between the specimen and Abelisauridae, Piatniktzysauridae, and Tyrannosauridae. MCT.R.514 was assigned as an abelisaurid due to its overall morphology that lacked metriacanthosaurid synapomorphies (i.e. spiraling mesial carina, strongly labially displaced distal carina, and well-marked mesiolingual longitudinal groove), while maintaining homoplastic features between both groups (i.e. irregular enamel texture, a lingually biconcave cross section, and a straight distal margin). This, together with a mostly Laurasian distribution of Metriacanthosauridae favors the assignment of MCT.R.514 as an abelisaurid as the most parsimonious hypothesis in this occasion. The identification of the specimen as an abelisaurid further expands the still scarce "Mid'' Cretaceous record of this clade in Brazil.
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Dinosaurios , Fósiles , Diente , Animales , Dinosaurios/clasificación , Dinosaurios/anatomía & histología , Brasil , Fósiles/anatomía & histología , Diente/anatomía & histología , FilogeniaRESUMEN
Despite making up one of the most ecologically diverse groups of living birds, comprising soaring, diving and giant flightless taxa, the evolutionary relationships and ecological evolution of Anseriformes (waterfowl) remain unresolved. Although Anseriformes have a comparatively rich, global Cretaceous and Paleogene fossil record, morphological datasets for this group that include extinct taxa report conflicting relationships for all known extinct taxa. Correct placement of extinct taxa is necessary to understand whether ancestral anseriform feeding ecology was more terrestrial or one of a set of diverse aquatic ecologies and to better understand avian evolution around the K-T boundary. Here, we present a new morphological dataset for Anseriformes that includes more extant and extinct taxa than any previous anseriform-focused dataset and describe a new anseriform species from the early Eocene Green River Formation of North America. The new taxon has a mediolaterally narrow bill which is rarely found in previously described anseriform fossils. The matrix created to assess the placement of this taxon comprises 41 taxa and 719 discrete morphological characters describing skeletal morphology, musculature, syringeal morphology, ecology, and behavior. We additionally combine the morphological dataset with published sequences using Bayesian methods and perform ancestral state reconstruction for select morphological, ecological and behavioral characters. We recover the new Eocene taxon as the sister taxon to (Anseranatidae+Anatidae) across all analyses, and find that the new taxon represents a novel ecology within known Anseriformes and the Green River taxa. Results provide insight into avian evolution during and following the K-Pg mass extinction and indicate that Anseriformes were likely ancestrally aquatic herbivores with rhamphothecal lamellae..
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Anseriformes , Evolución Biológica , Fósiles , Filogenia , Animales , Fósiles/anatomía & histología , Anseriformes/anatomía & histología , Anseriformes/clasificación , Anseriformes/genética , Aves/anatomía & histología , Aves/clasificaciónRESUMEN
The Karoo Basin of South Africa is renowned for its abundance and diversity of therapsid fossils. Among the most ubiquitous and persistent of the Permian fauna is the small herbivorous dicynodont Diictodon feliceps. Intraspecific variation in Diictodon is historically confounding, and while ontogeny is frequently cited as a potential source of variation, observable developmental changes have never been calibrated. The present study revisits this issue, comparing three-dimensional landmark configurations of 82 Diictodon crania to investigate the association between shape, size and dimorphism. Beyond the statistically significant relationship between shape and allometry, our results determine the shape differences between juvenile and adult skulls of Diictodon, aligned with common craniofacial features documented in other tetrapod taxa. Functionally, these changes are attributed to development of the jaw musculature for feeding on larger, tougher plant matter during later ontogeny. Cranial morphological variation owing to sexual dimorphism is negligible, but distinct differences are noted in the allometric trajectories of each morphotype. A component of non-allometric variation cannot be accounted for, and we propose that this represents natural variation, rather than an artefact of taphonomic deformation.
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Fósiles , Cráneo , Cráneo/anatomía & histología , Animales , Fósiles/anatomía & histología , Sudáfrica , Evolución Biológica , Dinosaurios/anatomía & histología , Dinosaurios/crecimiento & desarrollo , Caracteres Sexuales , MasculinoRESUMEN
Tyrannosaurids were the most derived group of Tyrannosauroidea and are characterized by having two body plans: gracile, long-snouted and robust, deep-snouted skulls. Both groups lived sympatrically in central Asia. Here, we report a new deep-snouted tyrannosaurid, Asiatyrannus xui gen. et sp. nov., from the Upper Cretaceous of Ganzhou City, southeastern China, which has produced the large-bodied and long-snouted Qianzhousaurus. Based on histological analysis, the holotype of Asiatyrannus xui is not a somatically mature adult, but it already passed through the most rapid growth stages. Asiatyrannus is a small to medium-sized tyrannosaurine, with a skull length of 47.5 cm and an estimated total body length of 3.5-4 m; or around half the size of Qianzhousaurus and other large-bodied tyrannosaurines in similar growth stages. Asiatyrannus and Qianzhousaurus are sympatric tyrannosaurid genera in the Maastrichtian of southeastern China. Asiatyrannus differs from Qianzhousaurus in that it has a proportionally deeper snout, longer premaxilla, deeper maxilla, and deeper dentary, and the cornual process of the lacrimal is inflated without developing a discrete horn. The different skull proportions and body sizes suggest that Asiatyrannus and Qianzhousaurus likely had different feeding strategies and occupied different ecological niches.
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Dinosaurios , Fósiles , Cráneo , Animales , Dinosaurios/anatomía & histología , Dinosaurios/clasificación , China , Fósiles/anatomía & histología , Cráneo/anatomía & histología , FilogeniaRESUMEN
A new troodontid dinosaur, Hypnovenator matsubaraetoheorum gen. et sp. nov., is described based on an articulated postcranial skeleton recovered from the fluvial deposits of the Albian Ohyamashimo Formation of the Sasayama Group in Tambasasayama City, Hyogo Prefecture, Japan. Hypnovenator is distinguished from other troodontids by four autapomorphies and a combination of additional features. Our phylogenetic analysis positions Hypnovenator as the oldest and one of the most basal troodontines, forming a clade with Gobivenator mongoliensis. The discovery of Hypnovenator suggests that small-bodied maniraptorans with a sleeping posture were common not only in environments with volcanic and eolian events or alluvial systems but also in fluvial systems. Geometric morphometric analysis of manual ungual phalanges shows that manual ungual phalanges I and III of Hypnovenator exhibit considerable morphological variation but are functionally similar, which differs from those of non-troodontine troodontids, reflecting the transition of manual motion within Troodontinae. Hypnovenator also has mosaic features in the pes related to cursoriality. This study reveals that asymmetrical arctometatarsus occurred by the Albian, and some morphological changes, such as shorter digit IV than digit III and non-ungual phalanges of digits III with roller joints and digit IV with weakly ginglymoid articulation, arose during the early Late Cretaceous.
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Evolución Biológica , Dinosaurios , Fósiles , Filogenia , Dinosaurios/anatomía & histología , Dinosaurios/clasificación , Animales , Fósiles/anatomía & histología , JapónRESUMEN
The diversity of cephalic morphologies in mandibulates (myriapods and pancrustaceans) was key to their evolutionary success. A group of Cambrian bivalved arthropods called hymenocarines exhibit diagnostic mandibulate traits that illustrate this diversity, but many forms are still poorly known. These include the odaraiids, typified by Odaraia alata from the Burgess Shale (Wuliuan), characterized by its unique tubular carapace and rudder-like tail fan, and one of the largest Cambrian euarthropods at nearly 20 cm in length. Unfortunately, odaraiid cephalic anatomy has been largely unknown, limiting evolutionary scenarios and putting their mandibulate affinities into question. Here, we reinvestigate Odaraia based on new specimens from the Burgess Shale and describe exquisitely preserved mandibles with teeth and adjacent structures: a hypostome, maxillae and potential paragnaths. These structures can be homologized with those of Cambrian fuxianhuiids and extant mandibulates, and suggest that the ancestral mandibulate head could have had a limbless segment but retained its plasticity, allowing for limb re-expression within Pancrustacea. Furthermore, we show the presence of limbs with spinose endites which created a suspension-feeding structure. This discovery provides morphological evidence for suspension feeding among large Cambrian euarthropods and evinces the increasing exploitation of planktonic resources in Cambrian pelagic food webs.
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Evolución Biológica , Fósiles , Animales , Fósiles/anatomía & histología , Artrópodos/anatomía & histología , Mandíbula/anatomía & histología , Conducta Alimentaria , FilogeniaRESUMEN
The announcement in 1925 by Raymond Dart of the discovery of the Taung juvenile's skull in a quarry in sub-Saharan Africa is deservedly a classic publication in the history of palaeoanthropology. Dart's paper-which designated Taung as the type specimen of the early hominin species Australopithecus africanus-provided the first fossil evidence supporting Charles Darwin's 1871 prediction that Africa was where the human lineage originated. The Taung juvenile's combination of ape and human characteristics eventually led to a paradigm shift in our understanding of human evolution. This contribution focuses on the milieu in which Dart's paper appeared (i.e. what was understood in 1925 about human evolution), the fossil evidence as set out by Dart, his interpretation of how a species represented by a fossilized juvenile's skull fitted within prevailing narratives about human evolution and the significance of the fossil being found in an environment inferred to be very different from that occupied by living apes. We also briefly review subsequent fossil finds that have corroborated the argument Dart made for having discovered evidence of a hitherto unknown close relative of humans, and summarize our current understanding of the earliest stages of human evolution and its environmental context.
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Evolución Biológica , Fósiles , Hominidae , Cráneo , Animales , Hominidae/anatomía & histología , Fósiles/anatomía & histología , Cráneo/anatomía & histología , Humanos , Historia del Siglo XX , ÁfricaRESUMEN
Recent studies suggest that both stem- and crown-group Archosauria encompassed high ecological diversity during their initial Triassic radiation. We describe a new pseudosuchian archosaur, Benggwigwishingasuchus eremicarminis gen. et sp. nov., from the Anisian (Middle Triassic) Fossil Hill Member of the Favret Formation (Nevada, USA), a pelagic setting in the eastern Panthalassan Ocean characterized by the presence of abundant ammonoids and large-bodied ichthyosaurs. Coupled with archosauriforms from the eastern and western Tethys Ocean, Benggwigwishingasuchus reveals that pseudosuchians were also components of Panthalassan ocean coastal settings, establishing that the group occupied these habitats globally during the Middle Triassic. However, Benggwigwishingasuchus, Qianosuchus, and Ticinosuchus (two other pseudosuchians known from marine sediments) are not recovered in a monophyletic group, demonstrating that a nearshore marine lifestyle occurred widely across Archosauriformes during this time. Benggwigwishingasuchus is recovered as part of an expanded Poposauroidea, including several taxa (e.g. Mandasuchus, Mambawakalae) from the Middle Triassic Manda Beds of Tanzania among its basally branching members. This implies a greater undiscovered diversity of poposauroids during the Early Triassic, and supports that the group, and pseudosuchians more broadly, diversified rapidly following the End-Permian mass extinction.
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Fósiles , Animales , Fósiles/anatomía & histología , Nevada , Filogenia , Reptiles/anatomía & histología , Reptiles/clasificación , Evolución Biológica , EcosistemaRESUMEN
Amber captures a snapshot of life and death from millions of years in the past. Here, the fate of three fossil Darwin wasps in Baltic amber is virtually dissected with the help of micro-CT scanning, to better understand the taphonomic processes that affected their preservation. The states of the fossils range from nearly perfect preservation, including remains of internal organs, to empty casts that were strongly affected by decomposition. We describe the three specimens as new taxa, Osparvis aurorae gen. et sp. nov., Grana harveydenti gen. et sp. nov. and Xorides? romeo sp. nov. Based on the taphonomic and morphological interpretations, we conclude that two specimens were trapped alive, and the third ended up in resin post-mortem. The morphology and classification of the specimens provide clues regarding their ecology, and we discuss their likely hosts and parasitation modes. Taken together, our three wasp fossils showcase how an integrative analysis of amber taphonomy, taxonomic association and morphology can shed light onto past biodiversity and offer valuable insights for interpreting their evolutionary history.
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Ámbar , Fósiles , Avispas , Animales , Fósiles/anatomía & histología , Avispas/anatomía & histología , Avispas/clasificación , Evolución Biológica , Microtomografía por Rayos XRESUMEN
Fragments of two skulls of young cetotheriid baleen whales were described from the Fortep'yanka 2 locality (Russia, Republic of Adygea, Maikop district, Fortep'yanka River valley, Upper Miocene, Upper Sarmatian, Blinovskaya Formation). The finds were attributed to Kurdalagonus maicopicus (Spasskii, 1951) based on the morphology of the posterior (mastoid) process of the petrosal bone, the structure of the posterior edge of the temporal fossa, and the S-like shape of the supraoccipital ridges. The skull proportions and the degree of suture closure made it possible to determine the individual age of the whales within a year. New finds significantly complement the data on the structure of the sutures of the lateral wall of the skull and age-related variability of cranial morphology in representatives of the genus Kurdalagonus.
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Fósiles , Cráneo , Animales , Cráneo/anatomía & histología , Fósiles/anatomía & histología , Federación de Rusia , Ballenas/anatomía & histología , Suturas Craneales/anatomía & histologíaRESUMEN
Pyritization of soft tissues of invertebrates is rare in the fossil record. In New York State, it occurs in black shales of the Lorraine Group (Late Ordovician), the best-known example of which is Beecher's Trilobite Bed. Exceptional preservation at the quarry where this bed is exposed allowed detailed examination of trilobite and ostracod soft-tissue anatomy. Here, we present the first example of a eurypterid (sea scorpion) currently ascribed to Carcinosomatidae from this deposit that also preserves the first evidence for mesosomal musculature in eurypterids. This specimen demonstrates that eurypterid musculature can be preserved in pyrite and evidences the oldest example of euchelicerate muscles within the fossil record. Sulfur isotope data illustrate that pyrite rapidly replicated muscle tissue in the early burial environment, prior to the pyritization of biomineralized exoskeleton and cuticular trilobite limbs. This discovery therefore expands the limited fossil record of euchelicerate musculature, while extending the taphonomic scope for preservation of detailed internal structures, more broadly, within arthropods.
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Fósiles , Hierro , Sulfuros , Animales , Fósiles/anatomía & histología , Músculos/anatomía & histología , Escorpiones/anatomía & histología , Isótopos de Azufre/análisis , New YorkRESUMEN
Accurate identification of waterfowl bones in archaeological and fossil assemblages has potential to unlock new methods of past environmental reconstruction, as species have differing habitat preferences and migration patterns. Therefore, identifying the presence of avian species with different ecological niches is key to determining past environments and ultimately how prehistoric people responded to climatic and environmental realignments. However, the identification of osteological remains of waterbirds such as ducks to species level is notoriously challenging. We address this by presenting a new two-dimensional geometric morphometric protocol on wing elements from over 20 duck species and test the utility of these shape data for correct species identification. This is an ideal starting point to expand utilization of these types of approaches in avifaunal research and test applicability to an extremely difficult taxonomic group.
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Patos , Alas de Animales , Animales , Alas de Animales/anatomía & histología , Patos/anatomía & histología , Osteología , Europa (Continente) , Fósiles/anatomía & histologíaRESUMEN
PREMISE: Characterization and phylogenetic integration of fossil angiosperms with uncertain affinities is relatively limited, which may obscure the diversity of extinct higher taxa in the flowering plant tree of life. The order Cornales contains a diversity of extinct taxa with uncertain familial affinities that make it an ideal group for studying turnover in angiosperms. Here, we describe a new extinct genus of Cornales unassignable to an extant family and conduct a series of phylogenetic analyses to reconstruct relationships of fossils across the order. METHODS: Two permineralized endocarps were collected from the Cedar District Formation (Campanian, 82-80 Ma) of Sucia Island, State of Washington, United States. Fossils were sectioned with the cellulose acetate peel technique and incorporated into a morphological dataset. To assess the utility of this dataset to accurately place taxa in their respective clades, we used a series of phylogenetic pseudofossilization analyses. We then conducted a total-evidence analysis and a scaffold-based approach to determine relationships of fossils. RESULTS: Based on their unique combination of characters, the fossils represent a new genus, Fenestracarpa washingtonensis gen. nov. et sp. nov. Pseudofossilization analyses indicate that our morphological dataset can be used to accurately recover taxa at the major clade to family level, generally with moderate to high support. The total-evidence and scaffold-based analyses recovered Fenestracarpa and other fossil genera in an entirely extinct clade within Cornales. CONCLUSIONS: Our findings increase the reported diversity of extinct Cornales and indicate that the order's initial radiation likely included the divergence of an extinct higher clade that endured the end-Cretaceous Mass extinction but perished during the Cenozoic.
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Extinción Biológica , Fósiles , Filogenia , Fósiles/anatomía & histología , Magnoliopsida/anatomía & histología , Magnoliopsida/genética , Magnoliopsida/clasificación , WashingtónAsunto(s)
Dieta , Fósiles , China , Fósiles/anatomía & histología , Animales , Desgaste de los Dientes , Ciervos , Diente/anatomía & histologíaRESUMEN
Scincidae is one of the most species-rich and cosmopolitan clades of squamate reptiles. Abundant disarticulated fossil material has also been attributed to this group, however, no complete pre-Cenozoic crown-scincid specimens have been found. A specimen in Burmite (99 MYA) is the first fossil that can be unambiguously referred to this clade. Our analyses place it as nested within extant skinks, supported by the presence of compound osteoderms formed by articulated small ostedermites. The specimen has a combination of dorsal and ventral compound osteoderms and overlapping cycloid scales that is limited to skinks. We propose that this type of osteoderm evolved as a response to an increased overlap of scales, and to reduced stiffness of the dermal armour. Compound osteoderms could be a key innovation that facilitated diversification in this megadiverse family.