RESUMEN
We redescribe the holotype and only known specimen of the early eureptile Coelostegus prothales from the Upper Carboniferous of the Czech Republic using photogrammetric scanning and a virtual 3D rendition of its skull. New information is available on several skull and lower jaw bones, including the postorbital, supratemporal, tabular, postparietal, angular, and prearticular. The new data also permit the correct identification of previously undetected or mis-identified elements (e.g., supratemporal; quadratojugal; angular). We provide an amended diagnosis of Coelostegus and a new reconstruction of the skull in dorsal and lateral views. To evaluate the affinities of Coelostegus, we code this taxon in two recently published taxon-character matrices. Parsimony and Bayesian analyses do not permit firm conclusions on the phylogenetic position of Coelostegus or, indeed, the status and extrinsic relationships of protorothyridid amniotes. Coelostegus emerges either as the sister taxon to the recently redefined Diapsida (Araeoscelidia; Varanopidae; Parareptilia; Neodiapsida), as one of the most basal protorothyridids, or as a derived stem-group amniote in various parsimony-based analyses, or as the basalmost protorothyridid in one Bayesian analysis, with protorothyridids forming a paraphyletic array relative to Diapsida. We review the cranial similarities and differences between Coelostegus and other protorothyridid genera and discuss the implications that various phylogenetic results have for our understanding of early amniote relationships.
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Cabeza , Cráneo , Teorema de Bayes , República Checa , Filogenia , AnimalesRESUMEN
Species richness is strikingly uneven across taxonomic groups at all hierarchical levels, but the reasons for this heterogeneity are poorly understood. It is well established that morphological diversity (disparity) is decoupled from taxonomic diversity, both between clades and across geological time. Morphological complexity has been much less studied, but there is theory linking complexity with differential diversity across groups. Here we devise an index of complexity from the differentiation of the fore and hind limb pairs for a sample of 983 species of extant birds. We test the null hypothesis that this index of morphological complexity is uncorrelated with clade diversity, revealing a significant and negative correlation between the species richness of clades and the mean morphological complexity of those clades. Further, we find that more complex clades tend to occupy a smaller number of dietary and habitat niches, and that this proxy for greater ecological specialisation correlates with lower species richness. Greater morphological complexity in the appendicular skeleton therefore appears to hinder the generation and maintenance of species diversity. This may result from entrenchment into morphologies and ecologies that are less capable of yielding further diversity.
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Aves , Geología , Animales , Miembro PosteriorRESUMEN
Avian blood parasites, from the genera Plasmodium, Haemoproteus and Leucocytozoon, are predicted to alter their range and prevalence as global temperatures change, and host and vector ranges shift. Understanding large-scale patterns in the prevalence and diversity of avian malaria and malaria-like parasites is important due to an incomplete understanding of their effects in the wild, where studies suggest even light parasitaemia can potentially cause rapid mortality, especially in naïve populations. We conducted phylogenetically controlled analyses to test for differences in prevalence and lineage diversity of haemoparasite infection (for Plasmodium, Haemoproteus and Leucocytozoon) in and between resident and migratory species along the African-Eurasian flyway. To test whether migratory strategy or range size drives differences in parasite prevalence and diversity between resident and migrant species, we included three categories of resident species: Eurasian only (n = 36 species), African only (n = 41), and species resident on both continents (n = 17), alongside intercontinental migrants (n = 64), using a subset of data from the MalAvi database comprising 27,861 individual birds. We found that species resident on both continents had a higher overall parasite diversity than all other categories. Eurasian residents had lower Plasmodium diversity than all other groups, and both migrants and species resident on both continents had higher Haemoproteus diversity than both African and Eurasian residents. Leucocytozoon diversity did not differ between groups. Prevalence patterns were less clear, with marked differences between genera. Both Plasmodium and Leucocytozoon prevalence was higher in species resident on both continents and African residents than in migrants and Eurasian residents. Haemoproteus prevalence was lower in Eurasian residents than species resident on both continents. Our findings contrast with previous findings in the North-South American flyway, where long-distance migrants had higher parasite diversity than residents and short-distance migrants, although we found contrasting patterns for parasite diversity to those seen for parasite prevalence. Crucially, our results suggest that geographic range may be more important than migratory strategy in driving parasite diversity within species along the African-Palaearctic flyway. Our findings differ between the three parasite genera included in our analysis, suggesting that vector ecology may be important in determining these large-scale patterns. Our results add to our understanding of global patterns in parasite diversity and abundance, and highlight the need to better understand the influence of vector ecology to understand the drivers of infection risk and predict responses to environmental change.
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Enfermedades de las Aves , Haemosporida , Parásitos , Plasmodium , Infecciones Protozoarias en Animales , Animales , Prevalencia , Enfermedades de las Aves/epidemiología , Enfermedades de las Aves/parasitología , Plasmodium/genética , Haemosporida/genética , Aves/parasitología , Filogenia , Infecciones Protozoarias en Animales/parasitologíaRESUMEN
Hostparasite associations provide a benchmark for investigating evolutionary arms races and antagonistic coevolution. However, potential ecological mechanisms underlying such associations are difficult to unravel. In particular, local adaptations of hosts and/or parasites may hamper reliable inferences of hostparasite relationships and the specialistgeneralist definitions of parasite lineages, making it problematic to understand such relationships on a global scale. Phylogenetic methods were used to investigate co-phylogenetic patterns between vector-borne parasites of the genus Haemoproteus and their passeriform hosts, to infer the ecological interactions of parasites and hosts that may have driven the evolution of both groups in a local geographic domain. As several Haemoproteus lineages were only detected once, and given the occurrence of a single extreme generalist, the effect of removing individual lineages on the co-phylogeny pattern was tested. When all lineages were included, and when all singly detected lineages were removed, there was no convincing evidence for hostparasite co-phylogeny. However, when only the generalist lineage was removed, strong support for co-phylogeny was indicated, and ecological interactions could be successfully inferred. This study exemplifies the importance of identifying locally abundant lineages when sampling hostparasite systems, to provide reliable insights into the precise mechanisms underlying hostparasite interactions.
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Enfermedades de las Aves , Haemosporida , Parásitos , Passeriformes , Animales , Filogenia , Haemosporida/genética , Interacciones Huésped-Parásitos , Enfermedades de las Aves/parasitologíaRESUMEN
Frogs exhibit complex anatomical features of the pelvis, limbs and spine, long assumed to represent specialisations for jumping. Yet frogs employ a wide range of locomotor modes, with several taxa featuring primary locomotor modes other than jumping. Using a combination of techniques (CT imaging and 3D visualization, morphometrics, phylogenetic mapping), this study aims to determine the link between skeletal anatomy and locomotor style, habitat type and phylogenetic history, shedding new light on how functional demands impact morphology. Body and limb measurements for 164 taxa from all the recognised anuran families are extracted from digitally segmented CT scans of whole frog skeletons and analysed using various statistical techniques. We find that the expansion of the sacral diapophyses is the most important variable for predicting locomotor mode, which was more closely correlated with frog morphology than either habitat type or phylogenetic relationships. Predictive analyses suggest that skeletal morphology is a useful indicator of jumping but less so for other locomotor modes, suggesting that there is a wide range of anatomical solutions to performing locomotor styles such as swimming, burrowing or walking.
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Evolución Biológica , Locomoción , Humanos , Animales , Filogenia , Anuros/anatomía & histología , NataciónRESUMEN
Complexity, defined as the number of parts and their degree of differentiation, is a poorly explored aspect of macroevolutionary dynamics. The maximum anatomical complexity of organisms has undoubtedly increased through evolutionary time. However, it is unclear whether this increase is a purely diffusive process or whether it is at least partly driven, occurring in parallel in most or many lineages and with increases in the minima as well as the means. Highly differentiated and serially repeated structures, such as vertebrae, are useful systems with which to investigate these patterns. We focus on the serial differentiation of the vertebral column in 1,136 extant mammal species, using two indices that quantify complexity as the numerical richness and proportional distribution of vertebrae across presacral regions and a third expressing the ratio between thoracic and lumbar vertebrae. We address three questions. First, we ask whether the distribution of complexity values in major mammal groups is similar or whether clades have specific signatures associated with their ecology. Second, we ask whether changes in complexity throughout the phylogeny are biased towards increases and whether there is evidence of driven trends. Third, we ask whether evolutionary shifts in complexity depart from a uniform Brownian motion model. Vertebral counts, but not complexity indices, differ significantly between major groups and exhibit greater within-group variation than recognized hitherto. We find strong evidence of a trend towards increasing complexity, where higher values propagate further increases in descendant lineages. Several increases are inferred to have coincided with major ecological or environmental shifts. We find support for multiple-rate models of evolution for all complexity metrics, suggesting that increases in complexity occurred in stepwise shifts, with evidence for widespread episodes of recent rapid divergence. Different subclades evolve more complex vertebral columns in different configurations and probably under different selective pressures and constraints, with widespread convergence on the same formulae. Further work should therefore focus on the ecological relevance of differences in complexity and a more detailed understanding of historical patterns.
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Evolución Biológica , Mamíferos , Animales , Filogenia , Columna VertebralRESUMEN
Phylogenetic relationships are inferred principally from two classes of data: morphological and molecular. Currently, most phylogenies of extant taxa are inferred from molecules and when morphological and molecular trees conflict the latter are often preferred. Although supported by simulations, the superiority of molecular trees has rarely been assessed empirically. Here we test phylogenetic accuracy using two independent data sources: biogeographic distributions and fossil first occurrences. For 48 pairs of morphological and molecular trees we show that, on average, molecular trees provide a better fit to biogeographic data than their morphological counterparts and that biogeographic congruence increases over research time. We find no significant differences in stratigraphic congruence between morphological and molecular trees. These results have implications for understanding the distribution of homoplasy in morphological data sets, the utility of morphology as a test of molecular hypotheses and the implications of analysing fossil groups for which molecular data are unavailable.
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Fósiles , FilogeniaRESUMEN
The taxonomically diverse terrestrial tetrapod fauna from the late Mississippian East Kirkton Limestone includes the earliest known members of stem Amphibia and stem Amniota. Here we name and describe a new East Kirkton tetrapod with an unusual hindlimb morphology reminiscent of that of several stem- and primitive crown amniotes. It displays a unique ilium with two slender and elongate processes and a 5-digit pes with a long, stout metatarsal IV and a greatly elongate digit IV. The new taxon broadens our knowledge of East Kirkton tetrapods, adding to the remarkable diversity of their hindlimb constructions, functional specializations, locomotory modes, and adaptations to a wide variety of substrates. An unweighted character parsimony analysis places the new taxon in a polytomy alongside some other Carboniferous groups. Conversely, weighted parsimony and Bayesian analyses retrieve it among the earliest diverging stem amniotes, either as the basalmost anthracosaur or within a clade that includes also Eldeceeon and Silvanerpeton, crownward of an array of chroniosaurs plus anthracosaurs.
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Evolución Biológica , Fósiles , Animales , Teorema de Bayes , Miembro Posterior , FilogeniaRESUMEN
Despite the increasing importance of molecular sequence data, morphology still makes an important contribution to resolving the phylogeny of many groups, and is the only source of data for most fossils. Most systematists sample morphological characters as broadly as possible on the principle of total evidence. However, it is not uncommon for sampling to be focused on particular aspects of anatomy, either because characters therein are believed to be more informative, or because preservation biases restrict what is available. Empirically, the optimal trees from partitions of morphological data sets often represent significantly different hypotheses of relationships. Previous work on hard-part versus soft-part characters across animal phyla revealed significant differences in about a half of sampled studies. Similarly, studies of the craniodental versus postcranial characters of vertebrates revealed significantly different trees in about one-third of cases, with the highest rates observed in non-avian dinosaurs. We test whether this is a generality here with a much larger sample of 81 published data matrices across all major dinosaur groups. Using the incongruence length difference test and two variants of the incongruence relationship difference test, we found significant incongruence in about 50% of cases. Incongruence is not uniformly distributed across major dinosaur clades, being highest (63%) in Theropoda and lowest (25%) in Thyreophora. As in previous studies, our partition tests show some sensitivity to matrix dimensions and the amount and distribution of missing entries. Levels of homoplasy and retained synapomorphy are similar between partitions, such that incongruence must partly reflect differences in patterns of homoplasy between partitions, which may itself be a function of modularity and mosaic evolution. Finally, we implement new tests to determine which partition yields trees most similar to those from the entire matrix. Despite no bias across dinosaurs overall, there are striking differences between major groups. The craniodental characters of Ornithischia and the postcranial characters of Saurischia yield trees most similar to the "total evidence" trees derived from the entire matrix. Trees from these same character partitions also tend to be most stratigraphically congruent: a mutual consilience suggesting that those partitions yield more accurate trees. [Dinosauria; homoplasy; partition homogeneity.].
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Dinosaurios/anatomía & histología , Dinosaurios/clasificación , Fósiles , Filogenia , Animales , Cráneo/anatomía & histología , Especificidad de la Especie , Diente/anatomía & histologíaRESUMEN
The known diversity of tetrapods of the Devonian period has increased markedly in recent decades, but their fossil record consists mostly of tantalizing fragments1-15. The framework for interpreting the morphology and palaeobiology of Devonian tetrapods is dominated by the near complete fossils of Ichthyostega and Acanthostega; the less complete, but partly reconstructable, Ventastega and Tulerpeton have supporting roles2,4,16-34. All four of these genera date to the late Famennian age (about 365-359 million years ago)-they are 10 million years younger than the earliest known tetrapod fragments5,10, and nearly 30 million years younger than the oldest known tetrapod footprints35. Here we describe Parmastega aelidae gen. et sp. nov., a tetrapod from Russia dated to the earliest Famennian age (about 372 million years ago), represented by three-dimensional material that enables the reconstruction of the skull and shoulder girdle. The raised orbits, lateral line canals and weakly ossified postcranial skeleton of P. aelidae suggest a largely aquatic, surface-cruising animal. In Bayesian and parsimony-based phylogenetic analyses, the majority of trees place Parmastega as a sister group to all other tetrapods.
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Fósiles , Filogenia , Vertebrados/anatomía & histología , Animales , Teorema de Bayes , Hueso Paladar/anatomía & histología , Federación de Rusia , Cráneo/anatomía & histología , Vertebrados/clasificaciónRESUMEN
Facial expression is a common channel for the communication of emotion. However, in the case of non-human animals, the analytical methods used to quantify facial expressions can be subjective, relying heavily on extrapolation from human-based systems. Here, we demonstrate how geometric morphometrics can be applied in order to overcome these problems. We used this approach to identify and quantify changes in facial shape associated with pain in a non-human animal species. Our method accommodates individual variability, species-specific facial anatomy, and postural effects. Facial images were captured at four different time points during ovariohysterectomy of domestic short haired cats (n = 29), with time points corresponding to varying intensities of pain. Images were annotated using landmarks specifically chosen for their relationship with underlying musculature, and relevance to cat-specific facial action units. Landmark data were subjected to normalisation before Principal Components (PCs) were extracted to identify key sources of facial shape variation, relative to pain intensity. A significant relationship between PC scores and a well-validated composite measure of post-operative pain in cats (UNESP-Botucatu MCPS tool) was evident, demonstrating good convergent validity between our geometric face model, and other metrics of pain detection. This study lays the foundation for the automatic, objective detection of emotional expressions in a range of non-human animal species.
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Cara/fisiología , Animales , Gatos , Emociones/fisiología , Expresión Facial , Humanos , Dimensión del Dolor , Dolor Postoperatorio/fisiopatologíaRESUMEN
The enigmatic tetrapod Acherontiscus caledoniae from the Pendleian stage of the Early Carboniferous shows heterodontous and durophagous teeth, representing the earliest known examples of significant adaptations in tetrapod dental morphology. Tetrapods of the Late Devonian and Early Carboniferous (Mississippian), now known in some depth, are generally conservative in their dentition and body morphologies. Their teeth are simple and uniform, being cone-like and sometimes recurved at the tip. Modifications such as keels occur for the first time in Early Carboniferous Tournaisian tetrapods. Acherontiscus, dated as from the Pendleian stage, is notable for being very small with a skull length of about 15 mm, having an elongate vertebral column and being limbless. Cladistic analysis places it close to the Early Carboniferous adelospondyls, aïstopods and colosteids and supports the hypothesis of 'lepospondyl' polyphyly. Heterodonty is associated with a varied diet in tetrapods, while durophagy suggests a diet that includes hard tissue such as chitin or shells. The mid-Carboniferous saw a significant increase in morphological innovation among tetrapods, with an expanded diversity of body forms, skull shapes and dentitions appearing for the first time.
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Adaptation is the fundamental driver of functional and biomechanical evolution. Accordingly, the states of biomechanical traits (absolute or relative trait values) have long been used as proxies for adaptations in response to direct selection. However, ignoring evolutionary history, in particular ancestry, passage of time and the rate of evolution, can be misleading. Here, we apply a recently developed phylogenetic statistical approach using significant rate shifts to detect instances of exceptional rates of adaptive changes in bite force in a large group of terrestrial vertebrates, the amniotes. Our results show that bite force in amniotes evolved through multiple bursts of exceptional rates of adaptive changes, whereby whole groups-including Darwin's finches, maniraptoran dinosaurs (group of non-avian dinosaurs including birds), anthropoids and hominins (fossil and modern humans)-experienced significant rate increases compared to the background rate. However, in most parts of the amniote tree of life, we find no exceptional rate increases, indicating that coevolution with body size was primarily responsible for the patterns observed in bite force. Our approach represents a template for future studies in functional morphology and biomechanics, where exceptional rates of adaptive changes can be quantified and potentially linked to specific ecological factors underpinning major evolutionary radiations.
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Adaptación Biológica , Evolución Biológica , Aves/fisiología , Mamíferos/fisiología , Reptiles/fisiología , Animales , Aves/anatomía & histología , Fuerza de la Mordida , Tamaño Corporal , Fósiles/anatomía & histología , Mamíferos/anatomía & histología , Filogenia , Reptiles/anatomía & histologíaRESUMEN
The pelvic girdle provides physical support and attachment for the hind limb musculature. In birds there is variability in pelvic morphology across different orders and this has been used as evidence for various types of locomotion. However, the morphological variation of pelvic bones has yet to be studied systematically in birds. Therefore, we investigated basic allometric relationships among female body mass (as a size proxy) and various pelvic measurements in a phylogenetic context. We also examined in detail the inter-relationships among various pelvic measurements. Also considered were the effects of broader taxonomic relationships at the level of order, with further examination of the influence of style of terrestrial locomotion on the allometric relationships. Positive relationships were initially found among all pelvic linear measurements and female body mass (FBM). The relationships among measures of pelvic width and FBM were isometric, whereas those between pelvic length and FBM showed positive allometry. Also, FBM explained more of the variance in pelvic length than in width. Similarly, the angle of the pelvis had a significant negative relationship, but FBM only explained a very low proportion of the variation in pelvic angles. In general terms, ancova showed that the effect of FBM was smaller than the effect of locomotor style in this species set. Both the synsacrum and pelvic girdle play roles in weight support and therefore scale in proportion to body weight accordingly. All three parts of the pelvis (ilium, ischium and pubis) are attached around the acetabulum and also provide muscle attachment points, so it might be expected for them to scale in a similar manner. Increased angulation of the pelvis is linked to orders which employ their hind limbs in feeding, perching and running, although FBM also explains a very low proportion of the variation in pelvic angle. Muscle attachment and the confines on morphology presented by locomotion explain much of the variation exhibited by the relationships among pelvic measurements.
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Aves/fisiología , Peso Corporal , Locomoción , Huesos Pélvicos/anatomía & histología , Filogenia , Animales , Biometría , Aves/anatomía & histología , Femenino , Análisis de Componente PrincipalRESUMEN
Phylogenetic tree shape varies as the evolutionary processes affecting a clade change over time. In this study, we examined an empirical phylogeny of fossil tetrapods during several time intervals, and studied how temporal constraints manifested in patterns of tree imbalance and character change. The results indicate that the impact of temporal constraints on tree shape is minimal and highlights the stability through time of the reference tetrapod phylogeny. Unexpected values of imbalance for Mississippian and Pennsylvanian time slices strongly support the hypothesis that the Carboniferous was a period of explosive tetrapod radiation. Several significant diversification shifts take place in the Mississippian and underpin increased terrestrialization among the earliest limbed vertebrates. Character incompatibility is relatively high at the beginning of tetrapod history, but quickly decreases to a relatively stable lower level, relative to a null distribution based on constant rates of character change. This implies that basal tetrapods had high, but declining, rates of homoplasy early in their evolutionary history, although the origin of Lissamphibia is an exception to this trend. The time slice approach is a powerful method of phylogenetic analysis and a useful tool for assessing the impact of combining extinct and extant taxa in phylogenetic analyses of large and speciose clades.
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Clasificación , Fósiles , Filogenia , Anfibios/clasificación , Animales , Evolución Biológica , Especiación GenéticaRESUMEN
Studies of the effects of mass extinctions on ancient ecosystems have focused on changes in taxic diversity, morphological disparity, abundance, behaviour and resource availability as key determinants of group survival. Crucially, the contribution of life history traits to survival during terrestrial mass extinctions has not been investigated, despite the critical role of such traits for population viability. We use bone microstructure and body size data to investigate the palaeoecological implications of changes in life history strategies in the therapsid forerunners of mammals before and after the Permo-Triassic Mass Extinction (PTME), the most catastrophic crisis in Phanerozoic history. Our results are consistent with truncated development, shortened life expectancies, elevated mortality rates and higher extinction risks amongst post-extinction species. Various simulations of ecological dynamics indicate that an earlier onset of reproduction leading to shortened generation times could explain the persistence of therapsids in the unpredictable, resource-limited Early Triassic environments, and help explain observed body size distributions of some disaster taxa (e.g., Lystrosaurus). Our study accounts for differential survival in mammal ancestors after the PTME and provides a methodological framework for quantifying survival strategies in other vertebrates during major biotic crises.
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Evolución Biológica , Huesos/anatomía & histología , Extinción Biológica , Fósiles , Animales , Tamaño Corporal , Cruzamiento , Dinosaurios , Planeta Tierra , Ecología , Ecosistema , Esperanza de Vida , Mamíferos , Dinámica Poblacional , Riesgo , VertebradosRESUMEN
The end-Devonian to mid-Mississippian time interval has long been known for its depauperate palaeontological record, especially for tetrapods. This interval encapsulates the time of increasing terrestriality among tetrapods, but only two Tournaisian localities previously produced tetrapod fossils. Here we describe five new Tournaisian tetrapods (Perittodus apsconditus, Koilops herma, Ossirarus kierani, Diploradus austiumensis and Aytonerpeton microps) from two localities in their environmental context. A phylogenetic analysis retrieved three taxa as stem tetrapods, interspersed among Devonian and Carboniferous forms, and two as stem amphibians, suggesting a deep split among crown tetrapods. We also illustrate new tetrapod specimens from these and additional localities in the Scottish Borders region. The new taxa and specimens suggest that tetrapod diversification was well established by the Tournaisian. Sedimentary evidence indicates that the tetrapod fossils are usually associated with sandy siltstones overlying wetland palaeosols. Tetrapods were probably living on vegetated surfaces that were subsequently flooded. We show that atmospheric oxygen levels were stable across the Devonian/Carboniferous boundary, and did not inhibit the evolution of terrestriality. This wealth of tetrapods from Tournaisian localities highlights the potential for discoveries elsewhere.
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Tree shape analyses are frequently used to infer the location of shifts in diversification rate within the Tree of Life. Many studies have supported a causal relationship between shifts and temporally coincident events such as the evolution of "key innovations". However, the evidence for such relationships is circumstantial. We investigated patterns of diversification during the early evolution of Amniota from the Carboniferous to the Triassic, subjecting a new supertree to analyses of tree balance in order to infer the timing and location of diversification shifts. We investigated how uneven origination and extinction rates drive diversification shifts, and use two case studies (herbivory and an aquatic lifestyle) to examine whether shifts tend to be contemporaneous with evolutionary novelties. Shifts within amniotes tend to occur during periods of elevated extinction, with mass extinctions coinciding with numerous and larger shifts. Diversification shifts occurring in clades that possess evolutionary innovations do not coincide temporally with the appearance of those innovations, but are instead deferred to periods of high extinction rate. We suggest such innovations did not cause increases in the rate of cladogenesis, but allowed clades to survive extinction events. We highlight the importance of examining general patterns of diversification before interpreting specific shifts.
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Organismos Acuáticos/clasificación , Extinción Biológica , Especiación Genética , Modelos Estadísticos , Filogenia , Amnios/fisiología , Animales , Organismos Acuáticos/genética , Biodiversidad , Fósiles , Herbivoria/genéticaRESUMEN
BACKGROUND: Adaptive radiation theory posits that ecological opportunity promotes rapid proliferation of phylogenetic and ecological diversity. Given that adaptive radiation proceeds via occupation of available niche space in newly accessed ecological zones, theory predicts that: (i) evolutionary diversification follows an 'early-burst' process, i.e., it accelerates early in the history of a clade (when available niche space facilitates speciation), and subsequently slows down as niche space becomes saturated by new species; and (ii) phylogenetic branching is accompanied by diversification of ecologically relevant phenotypic traits among newly evolving species. Here, we employ macroevolutionary phylogenetic model-selection analyses to address these two predictions about evolutionary diversification using one of the most exceptionally species-rich and ecologically diverse lineages of living vertebrates, the South American lizard genus Liolaemus. RESULTS: Our phylogenetic analyses lend support to a density-dependent lineage diversification model. However, the lineage through-time diversification curve does not provide strong support for an early burst. In contrast, the evolution of phenotypic (body size) relative disparity is high, significantly different from a Brownian model during approximately the last 5 million years of Liolaemus evolution. Model-fitting analyses also reject the 'early-burst' model of phenotypic evolution, and instead favour stabilizing selection (Ornstein-Uhlenbeck, with three peaks identified) as the best model for body size diversification. Finally, diversification rates tend to increase with smaller body size. CONCLUSIONS: Liolaemus have diversified under a density-dependent process with slightly pronounced apparent episodic pulses of lineage accumulation, which are compatible with the expected episodic ecological opportunity created by gradual uplifts of the Andes over the last ~25My. We argue that ecological opportunity can be strong and a crucial driver of adaptive radiations in continents, but may emerge less frequently (compared to islands) when major events (e.g., climatic, geographic) significantly modify environments. In contrast, body size diversification conforms to an Ornstein-Uhlenbeck model with multiple trait optima. Despite this asymmetric diversification between both lineages and phenotype, links are expected to exist between the two processes, as shown by our trait-dependent analyses of diversification. We finally suggest that the definition of adaptive radiation should not be conditioned by the existence of early-bursts of diversification, and should instead be generalized to lineages in which species and ecological diversity have evolved from a single ancestor.
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Lagartos/clasificación , Lagartos/genética , Animales , Evolución Biológica , Tamaño Corporal , Ecología , Especiación Genética , Modelos Genéticos , FilogeniaRESUMEN
Acanthostega is one of the earliest and most primitive limbed vertebrates. Its numerous fish-like features indicate a primarily aquatic lifestyle, yet cranial suture morphology suggests that its skull is more similar to those of terrestrial taxa. Here, we apply geometric morphometrics and two-dimensional finite-element analysis to the lower jaws of Acanthostega and 22 other tetrapodomorph taxa in order to quantify morphological and functional changes across the fish-tetrapod transition. The jaw of Acanthostega is similar to that of certain tetrapodomorph fish and transitional Devonian taxa both morphologically (as indicated by its proximity to those taxa in morphospace) and functionally (as indicated by the distribution of stress values and relative magnitude of bite force). Our results suggest a slow tempo of morphological and biomechanical changes in the transition from Devonian tetrapod jaws to aquatic/semi-aquatic Carboniferous tetrapod jaws. We conclude that Acanthostega retained a primitively aquatic lifestyle and did not possess cranial adaptations for terrestrial feeding.