Major Adaptive Radiations Research Articles

Stelladens mysteriosus: A Strange New Mosasaurid (Squamata) from the Maastrichtian (Late Cretaceous) of Morocco

Mosasaurids, a clade of specialized marine squamates, saw a major adaptive radiation in the Late Cretaceous, evolving a wide range of body sizes, shapes, and specialized tooth morphologies. The most diverse known mosasaurid faunas come from the late Maastrichtian phosphates of Morocco. Here, we report an unusual new mosasaurid, Stelladens mysteriosus, based on a partial jaw and associated tooth crowns from lower Couche III phosphatic deposits at Sidi Chennane, Oulad Abdoun Basin, Morocco. Stelladens is characterized by short, triangular tooth crowns with a series of strong, elaborate, and serrated ridges on the lingual surface of the tooth, functioning as accessory carinae. Morphology of the teeth and associated jaw fragment suggest affinities with Mosasaurinae. No close analogues to the unique tooth morphology of Stelladens are known, either extant or extinct. It may have had an unusual and highly specialized diet, a specialized prey-capture strategy, or both. The diversity of mosasaurid teeth is much higher than that of plesiosaurs, ichthyosaurs, or extant marine mammals, and likely reflects both the ecological diversity of mosasaurids and complex developmental mechanisms responsible for tooth formation in mosasaurines. Mosasaurid diversity continued to increase up to the Cretaceous–Paleogene boundary.

Recognising and quantifying the evolution of skeletal paedomorphosis in Plesiosauria

Plesiosaurs are one of the longest-ranging tetrapod groups in the Mesozoic and underwent a major adaptive radiation in the Late Triassic/Early Jurassic, so they are an ideal clade to study the long-term implications and deep-time evolution of specific developmental patterns. We compiled a database of all published plesiosaur specimens and recorded their skeletal maturity status. We use statistical modelling to demonstrate that the abundance of allegedly ‘juvenile’ specimens increases through time, which contradicts the null hypothesis that the relative proportion of juvenile to adult specimens should remain constant throughout evolution. These results indicate that many ‘juvenile’ specimens are really adults exhibiting heterochronic traits, particularly paedomorphism. Heterochrony is a developmental pattern particularly widespread in secondarily adapted organisms such as plesiosaurs. However, heterochronic patterns are typically only studied in individual genera/species or restricted clades. We demonstrate that the pervasiveness of paedomorphism in plesiosaurs increased gradually throughout the evolution of the clade, rather than being a specialization of specific clades.

Thermophilic Dehalococcoidia with unusual traits shed light on an unexpected past

Although the phylum Chloroflexota is ubiquitous, its biology and evolution are poorly understood due to limited cultivability. Here, we isolated two motile, thermophilic bacteria from hot spring sediments belonging to the genus Tepidiforma and class Dehalococcoidia within the phylum Chloroflexota. A combination of cryo-electron tomography, exometabolomics, and cultivation experiments using stable isotopes of carbon revealed three unusual traits: flagellar motility, a peptidoglycan-containing cell envelope, and heterotrophic activity on aromatics and plant-associated compounds. Outside of this genus, flagellar motility has not been observed in Chloroflexota, and peptidoglycan-containing cell envelopes have not been described in Dehalococcoidia. Although these traits are unusual among cultivated Chloroflexota and Dehalococcoidia, ancestral character state reconstructions showed flagellar motility and peptidoglycan-containing cell envelopes were ancestral within the Dehalococcoidia, and subsequently lost prior to a major adaptive radiation of Dehalococcoidia into marine environments. However, despite the predominantly vertical evolutionary histories of flagellar motility and peptidoglycan biosynthesis, the evolution of enzymes for degradation of aromatics and plant-associated compounds was predominantly horizontal and complex. Together, the presence of these unusual traits in Dehalococcoidia and their evolutionary histories raise new questions about the timing and selective forces driving their successful niche expansion into global oceans.

A subterranean adaptive radiation of amphipods in Europe

Adaptive radiations are bursts of evolutionary species diversification that have contributed to much of the species diversity on Earth. An exception is modern Europe, where descendants of ancient adaptive radiations went extinct, and extant adaptive radiations are small, recent and narrowly confined. However, not all legacy of old radiations has been lost. Subterranean environments, which are dark and food-deprived, yet buffered from climate change, have preserved ancient lineages. Here we provide evidence of an entirely subterranean adaptive radiation of the amphipod genus Niphargus, counting hundreds of species. Our modelling of lineage diversification and evolution of morphological and ecological traits using a time-calibrated multilocus phylogeny suggests a major adaptive radiation, comprised of multiple subordinate adaptive radiations. Their spatio-temporal origin coincides with the uplift of carbonate massifs in South-Eastern Europe 15 million years ago. Emerging subterranean environments likely provided unoccupied, predator-free space, constituting ecological opportunity, a key trigger of adaptive radiation. This discovery sheds new light on the biodiversity of Europe.

An early‐diverging procolophonid from the lowermost Triassic of South America and the origins of herbivory in Procolophonoidea

AbstractProcolophonoidea was the only parareptile clade to survive the end‐Permian mass extinction, and they experienced a major adaptive radiation during the Triassic. The flourishing of procolophonoid lineages in highly stressful post‐extinction environments may be, at least in part, a result of their early adaptations to herbivory, which would eventually become the most common feeding strategy in later representatives of the group. Although procolophonoids are comparatively diverse in the South African Lystrosaurus declivis Assemblage Zone, only the Gondwanan widespread taxon Procolophon trigoniceps has been thus far reported in South American deposits of similar age. Here, we present a new procolophonid genus and species from the Lower Triassic Sanga do Cabral Formation of Brazil. Oryporan insolitus gen. et sp. nov. combines typical adaptations suggestive of herbivory with primitive character states that are peculiar to early procolophonoids. Accordingly, phylogenetic analysis places the new species as an early‐diverging procolophonid, thus far the phylogenetically earliest to develop apparent adaptations for herbivory. Character optimization shows that the bulbous, labiolingually expanded molariforms of most procolophonids (usually interpreted as indicative of high‐fibre herbivory) were co‐opted from an insectivorous dentition. The new taxon also highlights the diversity of feeding habits displayed by Early Triassic procolophonoids, which was probably crucial for the survival of the clade in post‐extinction disaster communities.

Deep-Time Demographic Inference Suggests Ecological Release as Driver of Neoavian Adaptive Radiation

Assessing the applicability of theory to major adaptive radiations in deep time represents an extremely difficult problem in evolutionary biology. Neoaves, which includes 95% of living birds, is believed to have undergone a period of rapid diversification roughly coincident with the Cretaceous–Paleogene (K-Pg) boundary. We investigate whether basal neoavian lineages experienced an ecological release in response to ecological opportunity, as evidenced by density compensation. We estimated effective population sizes (Ne) of basal neoavian lineages by combining coalescent branch lengths (CBLs) and the numbers of generations between successive divergences. We used a modified version of Accurate Species TRee Algorithm (ASTRAL) to estimate CBLs directly from insertion–deletion (indel) data, as well as from gene trees using DNA sequence and/or indel data. We found that some divergences near the K-Pg boundary involved unexpectedly high gene tree discordance relative to the estimated number of generations between speciation events. The simplest explanation for this result is an increase in Ne, despite the caveats discussed herein. It appears that at least some early neoavian lineages, similar to the ancestor of the clade comprising doves, mesites, and sandgrouse, experienced ecological release near the time of the K-Pg mass extinction.

Phylogeny of Hawaiian Melicope (Rutaceae): RAD-seq Resolves Species Relationships and Reveals Ancient Introgression.

Hawaiian Melicope are one of the major adaptive radiations of the Hawaiian Islands comprising 54 endemic species. The lineage is monophyletic with an estimated crown age predating the rise of the current high islands. Phylogenetic inference based on Sanger sequencing has not been sufficient to resolve species or deeper level relationships. Here, we apply restriction site-associated DNA sequencing (RAD-seq) to the lineage to infer phylogenetic relationships. We employ Quartet Sampling to assess information content and statistical support, and to quantify discordance as well as partitioned ABBA-BABA tests to uncover evidence of introgression. Our new results drastically improved resolution of relationships within Hawaiian Melicope. The lineage is divided into five fully supported main clades, two of which correspond to morphologically circumscribed infrageneric groups. We provide evidence for both ancestral and current hybridization events. We confirm the necessity for a taxonomic revision of the Melicope section Pelea, as well as a re-evaluation of several species complexes by combining genomic and morphological data.

Taxonomic harmonization of Neogene and Quaternary candonid genera (Crustacea, Ostracoda) of the Paratethys

During the Cenozoic, the constant northward movement of the African plate led to the division of the Tethys Ocean into two: the Palaeomediterranean and the Paratethyan branches. The latter was represented by a huge epicontinental sea and brackish to freshwater lakes that extended across central Europe and western Asia. Neogene and Quaternary ostracods from the Paratethys originated through major adaptive radiations, which gave rise to endemic brackish taxa. Unfortunately, much confusion surrounds their taxonomy, due to the quality of descriptions and images in the original literature and the incompleteness of the type material, making necessary a taxonomic revision. In this paper, we propose a systematic revision of several Paratethyan endemic candonid genera based on the analysis of the type material, new material collected from the type localities, and new fossil material from the Ponto-Caspian area. The study focuses on the description of the valve morphology and particularly on the geometric morphometric analysis of the valve outline. Thirty-three genera were taken into account of which four (Advenocypris, Candoniella, Graviacypris, Telekia) were considered to be junior synonyms of Typhlocypris, Pseudocandona or Candona. Moesiella is considered a nomen nudum. In the case of Caspiollina, Dacicandona, Liventalina and Turkmenella the scarcity of material and/or the poor descriptions reported in the literature prevented us from performing a full revision. The monospecific genus Thaminocypris possibly includes a teratological form. The remaining 23 genera were merged into nine valid, endemic, genera (Bakunella, Camptocypria, Caspiocypris, Hastacandona, Lineocypris, Pontoniella, Propontoniella, Typhlocyprella and Zalanyiella). Emended diagnoses and descriptions are proposed for these genera and a new species, Bakunella anae sp. nov., is described. This study considerably reduces the taxonomic uncertainty within the Paratethyan candonids, providing new data for the evaluation of the palaeobiodiversity of the Paratethyan domain. Finally, the palaeobiogeography of Paratethyan candonids during the Neogene and Quaternary is also discussed.http://zoobank.org/urn:lsid:zoobank.org:pub:EB9EC2D4-AFD2-428E-9958-C97F36ED7FF2

A Jurassic gliding euharamiyidan mammal with an ear of five auditory bones.

Gliding is a distinctive locomotion type that has been identified in only three mammal species from the Mesozoic era. Here we describe another Jurassic glider that belongs to the euharamiyidan mammals and shows hair details on its gliding membrane that are highly similar to those of extant gliding mammals. This species possesses a five-boned auditory apparatus consisting of the stapes, incus, malleus, ectotympanic and surangular, representing, to our knowledge, the earliest known definitive mammalian middle ear. The surangular has not been previously identified in any mammalian middle ear, and the morphology of each auditory bone differs from those of known mammals and their kin. We conclude that gliding locomotion was probably common in euharamiyidans, which lends support to idea that there was a major adaptive radiation of mammals in the mid-Jurassic period. The acquisition of the auditory bones in euharamiyidans was related to the formation of the dentary-squamosal jaw joint, which allows a posterior chewing movement, and must have evolved independently from the middle ear structures of monotremes and therian mammals.

Polyploidy and interspecific hybridization: partners for adaptation, speciation and evolution in plants.

Polyploidy or whole-genome duplication is now recognized as being present in almost all lineages of higher plants, with multiple rounds of polyploidy occurring in most extant species. The ancient evolutionary events have been identified through genome sequence analysis, while recent hybridization events are found in about half of the world's crops and wild species. Building from this new paradigm for understanding plant evolution, the papers in this Special Issue address questions about polyploidy in ecology, adaptation, reproduction and speciation of wild and cultivated plants from diverse ecosystems. Other papers, including this review, consider genomic aspects of polyploidy. Discovery of the evolutionary consequences of new, evolutionarily recent and ancient polyploidy requires a range of approaches. Large-scale studies of both single species and whole ecosystems, with hundreds to tens of thousands of individuals, sometimes involving 'garden' or transplant experiments, are important for studying adaptation. Molecular studies of genomes are needed to measure diversity in genotypes, showing ancestors, the nature and number of polyploidy and backcross events that have occurred, and allowing analysis of gene expression and transposable element activation. Speciation events and the impact of reticulate evolution require comprehensive phylogenetic analyses and can be assisted by resynthesis of hybrids. In this Special Issue, we include studies ranging in scope from experimental and genomic, through ecological to more theoretical. The success of polyploidy, displacing the diploid ancestors of almost all plants, is well illustrated by the huge angiosperm diversity that is assumed to originate from recurrent polyploidization events. Strikingly, polyploidization often occurred prior to or simultaneously with major evolutionary transitions and adaptive radiation of species, supporting the concept that polyploidy plays a predominant role in bursts of adaptive speciation. Polyploidy results in immediate genetic redundancy and represents, with the emergence of new gene functions, an important source of novelty. Along with recombination, gene mutation, transposon activity and chromosomal rearrangement, polyploidy and whole-genome duplication act as drivers of evolution and divergence in plant behaviour and gene function, enabling diversification, speciation and hence plant evolution.

The tarsal-metatarsal complex of caviomorph rodents: Anatomy and functional-adaptive analysis.

Caviomorph rodents represent a major adaptive radiation of Neotropical mammals. They occupy a variety of ecological niches, which is also reflected in their wide array of locomotor behaviors. It is expected that this radiation would be mirrored by an equivalent disparity of tarsal-metatarsal morphology. Here, the tarsal-metatarsal complex of Erethizontidae, Cuniculidae, Dasyproctidae, Caviidae, Chinchillidae, Octodontidae, Ctenomyidae, and Echimyidae was examined, in order to evaluate its anatomical variation and functional-adaptive relevance in relation to locomotor behaviors. A qualitative study in functional morphology and a geometric morphometric analysis were performed. We recognized two distinct tarsal-metatarsal patterns that represent the extremes of anatomical variation in the foot. The first, typically present in arboreal species, is characterized by features that facilitate movements at different levels of the tarsal-metatarsal complex. The second pattern, typically present in cursorial caviomorphs, has a set of features that act to stabilize the joints, improve the interlocking of the tarsal bones, and restrict movements to the parasagittal plane. The morphological disparity recognized in this study seems to result from specific locomotor adaptations to climb, dig, run, jump and swim, as well as phylogenetic effects within and among the groups studies.

Gene Expression Variation Resolves Species and Individual Strains among Coral-Associated Dinoflagellates within the Genus Symbiodinium.

Reef-building corals depend on symbiotic mutualisms with photosynthetic dinoflagellates in the genus Symbiodinium. This large microalgal group comprises many highly divergent lineages (“Clades A–I”) and hundreds of undescribed species. Given their ecological importance, efforts have turned to genomic approaches to characterize the functional ecology of Symbiodinium. To date, investigators have only compared gene expression between representatives from separate clades—the equivalent of contrasting genera or families in other dinoflagellate groups—making it impossible to distinguish between clade-level and species-level functional differences. Here, we examined the transcriptomes of four species within one Symbiodinium clade (Clade B) at ∼20,000 orthologous genes, as well as multiple isoclonal cell lines within species (i.e., cultured strains). These species span two major adaptive radiations within Clade B, each encompassing both host-specialized and ecologically cryptic taxa. Species-specific expression differences were consistently enriched for photosynthesis-related genes, likely reflecting selection pressures driving niche diversification. Transcriptional variation among strains involved fatty acid metabolism and biosynthesis pathways. Such differences among individuals are potentially a major source of physiological variation, contributing to the functional diversity of coral holobionts composed of unique host–symbiont genotype pairings. Our findings expand the genomic resources available for this important symbiont group and emphasize the power of comparative transcriptomics as a method for studying speciation processes and interindividual variation in nonmodel organisms.

Evidence for a Mid-Jurassic Adaptive Radiation in Mammals

A series of spectacular discoveries have transformed our understanding of Mesozoic mammals in recent years. These finds reveal hitherto-unsuspected ecomorphological diversity that suggests that mammals experienced a major adaptive radiation during the Middle to Late Jurassic. Patterns of mammalian macroevolution must be reinterpreted in light of these new discoveries, but only taxonomic diversity and limited aspects of morphological disparity have been quantified. We assess rates of morphological evolution and temporal patterns of disparity using large datasets of discrete characters. Rates of morphological evolution were significantly elevated prior to the Late Jurassic, with a pronounced peak occurring during the Early to Middle Jurassic. This intense burst of phenotypic innovation coincided with a stepwise increase in apparent long-term standing diversity and the attainment of maximum disparity, supporting a "short-fuse" model of early mammalian diversification. Rates then declined sharply, and remained significantly low until the end of the Mesozoic, even among therians. This supports the "long-fuse" model of diversification in Mesozoic therians. Our findings demonstrate that sustained morphological innovation in Triassic stem-group mammals culminated in a global adaptive radiation of crown-group members during the Early to Middle Jurassic.

Phylogeny, Ecology, and Morphological Evolution in the Atelid Cranium

Reconstructing evolutionary relationships of living and extinct primate groups requires reliable phylogenetic inference based on morphology, as DNA is rarely preserved in fossil specimens. Atelids (family Atelidae) are a monophyletic clade and one of the three major adaptive radiations of south and central American primates (platyrrhines), including the genera Alouatta, Ateles, Brachyteles, and Lagothrix, and are diverse in morphology, body and brain size, locomotion, diet, social systems, and behavioral ecology. Molecular phylogenetic relationships of the extant atelid genera are well resolved, yet morphological analyses often support alternative phylogenetic relationships to molecular data. We collected geometric morphometric data from the crania of atelid taxa for phylogenetic analysis of the cranium, cranial base, and face and tested the hypotheses that cranial data maintain a phylogenetic signal, cranial base morphology most closely reflects the atelid molecular phylogeny, and facial and overall cranial morphology are shaped by diet and have experienced greater homoplasy. All analyses supported genus monophyly, and facial morphology maintained a strong phylogenetic signal inferring the atelid molecular phylogeny and a sister relationship between Brachyteles and Lagothrix, whereas results from the cranial base and whole cranium supported Ateles–Lagothrix and/or Alouatta–Brachyteles clades reflecting homoplasy and ecological and dietary similarities. A phylogenetic signal in the atelid face is important for future studies integrating fossil taxa and supports evidence that congruence between molecular and morphological phylogenetics in primates is module and clade specific.

Hybrid breakdown in cichlid fish.

Studies from a wide diversity of taxa have shown a negative relationship between genetic compatibility and the divergence time of hybridizing genomes. Theory predicts the main breakdown of fitness to happen after the F1 hybrid generation, when heterosis subsides and recessive allelic (Dobzhansky-Muller) incompatibilities are increasingly unmasked. We measured the fitness of F2 hybrids of African haplochromine cichlid fish bred from species pairs spanning several thousand to several million years divergence time. F2 hybrids consistently showed the lowest viability compared to F1 hybrids and non-hybrid crosses (crosses within the grandparental species), in agreement with hybrid breakdown. Especially the short- and long-term survival (2 weeks to 6 months) of F2 hybrids was significantly reduced. Overall, F2 hybrids showed a fitness reduction of 21% compared to F1 hybrids, and a reduction of 43% compared to the grandparental, non-hybrid crosses. We further observed a decrease of F2 hybrid viability with the genetic distance between grandparental lineages, suggesting an important role for negative epistatic interactions in cichlid fish postzygotic isolation. The estimated time window for successful production of F2 hybrids resulting from our data is consistent with the estimated divergence time between the multiple ancestral lineages that presumably hybridized in three major adaptive radiations of African cichlids.

Evolutionary diversification in freshwater sculpins (Cottoidea): a review of two major adaptive radiations

Freshwater sculpins, grouped by their common ecological characteristic of freshwater habitats, comprise the species from the genera Cottus, Trachidermus, Mesocottus and Myoxocephalus as well as the Baikal sculpins. These fishes are typically coldwater-adapted, having probably originated polyphyletically from ancestral species of marine sculpins. Gottus, the most speciose taxonomic group, includes at least 64 species and is distributed throughout the fresh waters of the northern part of the Northern Hemisphere. Indinidual species have diverse life histories, such as fluvial, lacustrine, catadromous, and freshwater amphidromous. The second most abundant group, the Baikal sculpins, includes 33 species in 12 genera representing 3 families, and comprises many benthic, and a few bentho-pelagic and pelagic species. The freshwater sculpins belonging to Trachidermus, Mesocottus and Myoxocephalus include only one or two species in each genus. Recent molecular phylogenetic analyses of Cottus species, Baikal sculpins and their relatives have demonstrated mainly that (1) Cottus kazika is a sister taxon to Trachidermus fasciatus (designated lineage A), (2) Cottus species, except for C. kazika, and the Baikal sculpins are monophyletic, (3) on the basis of (1) and (2), Cottus as presently recognized is not monophyletic, and (4) the Major monophyletic lineages include 7 lineages: lineage B from Eurasia, lineages C and D from East Asia, the Cottopsis clade (sensu Copeia 2005:303–311, 2005) from the west coast of North America, the Cottus clade from the Circum-Arctic sea, the monophyletic Baikal sculpins, and the Uranidea clade. These findings suggested that the monophyletic freshwater sculpins that comprise the lineage A and the 7 other lineages may undergone two major radiations, one having occurred in the fresh water Cottus species in the northern part of the Northern Hemisphere, and the other in the Baikal sculpins in the Lake Baikal, the world’s oldest freshwater lake. Through these adaptive radiations, a tremendous diversity of morphological, ecological, physiological and life historical traits now exists in the freshwater sculpins.

Species compositions and ecology of the riverine ichthyofaunas in two Sulawesian islands in the biodiversity hotspot of Wallacea

This account of the riverine ichthyofaunas from the islands of Buton and Kabaena, off south-eastern mainland Sulawesi, represents the first detailed quantitative checklist and ecological study of the riverine fish faunas in the biological hotspot of Wallacea. The results are based on analysis of samples collected by electrofishing at a wide range of sites from July to September in both 2001 and 2002. While the fauna was diverse, with the 2179 fishes caught comprising 64 species representing 43 genera and 22 families, the catches were dominated by the Gobiidae (26 species and 25% by numbers), Eleotridae (seven species and 27% by numbers), Zenarchopteridae (three species and 22% by numbers) and Anguillidae (two species and 12% by numbers). The most abundant species were the eleotrids Eleotris aff. fusca-melanosoma and Ophieleotris aff. aporos, the anguillid Anguilla celebesensis, the zenarchopterids Nomorhamphus sp. and Nomorhamphus ebrardtii and the gobiids Sicyopterus sp. and Glossogobius aff. celebius-kokius. The introduced catfish Clarias batrachus was moderately abundant at a few sites. Cluster analysis, allied with the similarity profiles routine SIMPROF, identified seven discrete groups, which represented samples from sites entirely or predominantly in either Buton (five clusters) or Kabaena (two clusters). Species composition was related to geographical location, distance from river mouth, per cent contribution of sand and silt, altitude and water temperature. The samples from the two islands contained only one species definitively endemic to Sulawesi, i.e. N. ebrardtii and another presumably so, i.e. Nomorhamphus sp., contrasting starkly with the 57 species that are endemic to Sulawesi and, most notably, its large central and deep lake systems on the mainland. This accounts for the ichthyofaunas of these two islands, as well as those of rivers in northern mainland Sulawesi and Flores, being more similar to each other than to those of the central mainland lake systems. This implies that the major adaptive radiation of freshwater fishes in Sulawesi occurred in those lacustrine environments rather than in rivers.

Late to the Table: Diversification of Tetrapod Mandibular Biomechanics Lagged Behind the Evolution of Terrestriality

The origin of digit-bearing tetrapods in the Middle to Late Devonian (ca. 393-360 Mya) and their subsequent invasion of land represent a classic case of a major evolutionary radiation driven by new ecological opportunities. In this and other examples, exploration of new eco-space is hypothesized to correlate with functional innovation and adaptive divergence of phenotypes. Simultaneous changes in all morphofunctional systems are rare in major evolutionary transitions and may be non-existent. Here, we focus on the mandibles of early tetrapods and their kin as a model system to test whether shifts in functional innovation were coeval with some major events in tetrapod history. To this end, we quantified mechanical variation in the mandibles of tetrapodomorphs ranging in age from Early Devonian to earliest Permian. Biomechanical disparity is stable from the Devonian to the Early Pennsylvanian, even though the origin of weight-bearing, digited limbs, and the initial phases of the colonization of land occurred during this interval. An appreciable increase in functional variation is detected in the latest Pennsylvanian and earliest Permian, when stem and crown amniotes began to explore new regions of mechanical morphospace, a pattern partly attributable to the origin of herbivory. We find no difference in the rate of functional change between tetrapodomorph "fish" and early digited tetrapods, although two independent shifts are detected among Devonian stem tetrapods more crownward than Acanthostega. Instead, the most profound shifts in evolutionary rate are nested well within the tetrapod crown and are associated with amniotes (particularly diadectomorphs and some synapsids). The substantial temporal gap between the origin of postcranial features associated with terrestriality, such as limbs with functional elbow/knee and wrist/ankle joints capable of weight-bearing, and the onset of divergence in jaw biomechanics provides a compelling example of "functional modularity" during a major adaptive radiation.

Primates

Primates

A transitional snake from the Late Cretaceous period of North America

Snakes are the most diverse group of lizards, but their origins and early evolution remain poorly understood owing to a lack of transitional forms. Several major issues remain outstanding, such as whether snakes originated in a marine or terrestrial environment and how their unique feeding mechanism evolved. The Cretaceous Coniophis precedens was among the first Mesozoic snakes discovered, but until now only an isolated vertebra has been described and it has therefore been overlooked in discussions of snake evolution. Here we report on previously undescribed material from this ancient snake, including the maxilla, dentary and additional vertebrae. Coniophis is not an anilioid as previously thought a revised phylogenetic analysis of Ophidia shows that it instead represents the most primitive known snake. Accordingly, its morphology and ecology are critical to understanding snake evolution. Coniophis occurs in a continental floodplain environment, consistent with a terrestrial rather than a marine origin; furthermore, its small size and reduced neural spines indicate fossorial habits, suggesting that snakes evolved from burrowing lizards. The skull is intermediate between that of lizards and snakes. Hooked teeth and an intramandibular joint indicate that Coniophis fed on relatively large, soft-bodied prey. However, the maxilla is firmly united with the skull, indicating an akinetic rostrum. Coniophis therefore represents a transitional snake, combining a snake-like body and a lizard-like head. Subsequent to the evolution of a serpentine body and carnivory, snakes evolved a highly specialized, kinetic skull, which was followed by a major adaptive radiation in the Early Cretaceous period. This pattern suggests that the kinetic skull was a key innovation that permitted the diversification of snakes.