Macroevolutionary Trends Research Articles

Cautionary tales on the use of proxies to estimate body size and form of extinct animals.

Body size is of fundamental importance to our understanding of extinct organisms. Physiology, ecology and life history are all strongly influenced by body size and shape, which ultimately determine how a species interacts with its environment. Reconstruction of body size and form in extinct animals provides insight into the dynamics underlying community composition and faunal turnover in past ecosystems and broad macroevolutionary trends. Many extinct animals are known only from incomplete remains, necessitating the use of anatomical proxies to reconstruct body size and form. Numerous limitations affecting the appropriateness of these proxies are often overlooked, leading to controversy and downstream inaccuracies in studies for which reconstructions represent key input data. In this perspective, we discuss four prominent case studies (Dunkleosteus, Helicoprion, Megalodon and Perucetus) in which proxy taxa have been used to estimate body size and shape from fragmentary remains. We synthesise the results of these and other studies to discuss nuances affecting the validity of taxon selection when reconstructing extinct organisms, as well as mitigation measures that can ensure the selection of the most appropriate proxy. We argue that these precautionary measures are necessary to maximise the robustness of reconstructions in extinct taxa for better evolutionary and ecological inferences.

Understanding niche construction and phenotypic plasticity as causes of natural selection

AbstractFor many evolutionary biologists, fitness differences cause trait frequency changes in populations, and natural selection explains the evolution of adaptations. Treating fitness differences as a cause, however, is more scientific convention rather than decree, and analyses of the causes of natural selection potentially afford richer evolutionary explanations. Unfortunately, the historical assumptions that the complexities of development leave the origins of phenotypic variation unpredictable, and that ecological processes are idiosyncratic, have hindered detailed analysis of the developmental bases of natural selection. A poorly appreciated consequence is that explanations reliant on selection potentially mask particular causal patterns important in evolution. Here, using examples of environmental modification and regulation by organisms (‘niche construction’, a.k.a. ‘ecosystem engineering’), and developmental plastic responses to environmental conditions (‘phenotypic plasticity’), I will highlight how the development and activities of organisms create developmental biases that co‐determine the nature of the response to selection, in an often surprisingly well‐regulated manner. Niche construction biases the phenotypic variation exposed to selection, often generating axes of covariation with plastically expressed morphological traits. Taxonomically shared developmental mechanisms aggregate across populations to generate statistical regularities that are easy to miss because the developmental causes of fitness differences are not currently central to the study of evolution. Recent theory and experiments suggest that how organisms develop and what organisms do cause and strengthen the relationship between key traits and fitness, thereby part‐determining the characteristics of natural selection. The findings have implications for understanding parallel evolution, macroevolutionary trends and variation in evolvability.

Comparative functional morphology indicates niche partitioning among sympatric marine reptiles.

Mesozoic marine ecosystems were dominated by diverse lineages of aquatic tetrapods. For over 50 Ma in the Jurassic until the Early Cretaceous, plesiosaurians, ichthyosaurians and thalattosuchian crocodylomorphs coexisted at the top levels of trophic food webs. We created a functional dataset of continuous craniomandibular and dental characters known from neontological studies to be functionally significant in modern aquatic tetrapods. We analysed this dataset with multivariate ordination and inferential statistics to assess functional similarities and differences in the marine reptile faunas of two well-sampled Jurassic ecosystems deposited in the same seaway: the Oxford Clay Formation (OCF, Callovian-early Oxfordian, Middle-Late Jurassic) and the Kimmeridge Clay Formation (KCF, Kimmeridgian-Tithonian, Late Jurassic) of the UK. Lower jaw-based macroevolutionary trends are similar to those of tooth-based diversity studies. Closely related species cluster together, with minimal overlaps in the morphospace. Marine reptile lineages were characterized by the distinctive combinations of features, but we reveal multiple instances of morphofunctional convergence among different groups. We quantitatively corroborate previous observations that the ecosystems in the OCF and KCF were markedly distinct in faunal composition and structure. Morphofunctional differentiation may have enabled specialization and was an important factor facilitating the coexistence of diverse marine reptile assemblages in deep time.

Varying Modes of Selection Among Toxin Families in the Venoms of the Giant Desert Hairy Scorpions (Hadrurus).

Venoms are primarily believed to evolve under strong diversifying selection resulting from persistent coevolution between predator and prey. Recent research has challenged this hypothesis, proposing that venoms from younger venomous lineages (e.g., snakes and cone snails) are governed predominantly by diversifying selection, while venoms from older venomous lineages (e.g., centipedes, scorpions, and spiders) are under stronger purifying selection. However, most research in older lineages has tested selection at more diverse phylogenetic scales. Although these tests are important for evaluating broad macroevolutionary trends underlying venom evolution, they are less equipped to detect species-level evolutionary trends, which likely have large impacts on venom variation seen at more diverse phylogenetic scales. To test for selection among closely related species from an older venomous lineage, we generated high-throughput venom-gland transcriptomes and venom proteomes for four populations of Giant Desert Hairy Scorpions (Hadrurus), including three Hadrurus arizonensis populations and one Hadrurus spadix population. We detected significant episodic and pervasive diversifying selection across a highly abundant toxin family that likely has a major role in venom function ([Formula: see text]KTxs), providing a contrast to the stronger purifying selection identified from other studies on scorpion venoms. Conversely, we detected weak episodic diversifying and/or stronger purifying selection in four toxin families (non-disulfide bridged peptides, phospholipase A2s, scorpine-like antimicrobial peptides, and serine proteases), most of which were less abundant and likely have ancillary functional roles. Finally, although we detected several major toxin families at disproportionate transcriptomic and/or proteomic abundances, we did not identify significant sex-based variation in Hadrurus venoms.

The neck as a keystone structure in avian macroevolution and mosaicism

BackgroundThe origin of birds from non-avian theropod dinosaur ancestors required a comprehensive restructuring of the body plan to enable the evolution of powered flight. One of the proposed key mechanisms that allowed birds to acquire flight and modify the associated anatomical structures into diverse forms is mosaic evolution, which describes the parcelization of phenotypic traits into separate modules that evolve with heterogeneous tempo and mode. Avian mosaicism has been investigated with a focus on the cranial and appendicular skeleton, and as such, we do not understand the role of the axial column in avian macroevolution. The long, flexible neck of extant birds lies between the cranial and pectoral modules and represents an opportunity to study the contribution of the axial skeleton to avian mosaicism.ResultsHere, we use 3D geometric morphometrics in tandem with phylogenetic comparative methods to provide, to our knowledge, the first integrative analysis of avian neck evolution in context with the head and wing and to interrogate how the interactions between these anatomical systems have influenced macroevolutionary trends across a broad sample of extant birds. We find that the neck is integrated with both the head and the forelimb. These patterns of integration are variable across clades, and only specific ecological groups exhibit either head-neck or neck-forelimb integration. Finally, we find that ecological groups that display head-neck and neck-forelimb integration tend to display significant shifts in the rate of neck morphological evolution.ConclusionsCombined, these results suggest that the interaction between trophic ecology and head-neck-forelimb mosaicism influences the evolutionary variance of the avian neck. By linking together the biomechanical functions of these distinct anatomical systems, the cervical vertebral column serves as a keystone structure in avian mosaicism and macroevolution.

Scolicia, ichnotaxonomic practices, and the limits of behavioural convergence

Many studies convincingly documented the diagnostic morphology of Scolicia, establishing a functional link with the burrowing activities of heart urchins, common components of the marine mobile faunas that emerged during the Mesozoic Marine Revolution. However, this ichnotaxon has been repeatedly mentioned in both continental successions and Paleozoic marine strata, even though its diagnostic characteristics were absent. This approach may reflect the implicit belief that behavioural convergence is rampant in the ichnologic record and can be uncritically adopted in almost all cases. We underscore the importance of a definition of Scolicia that is based on diagnostic morphologic features and supported by a solid analysis from the perspective of function and mode of construction. This view of Scolicia highlights its value in paleoenvironmental reconstructions and its significance to unravel macroevolutionary trends. Attribution to an ichnotaxon should involve understanding of morphologic features in terms of its mode of production rather than the mere observation of superficial similarities.

Dating the evolution of the complex thalloid liverworts (Marchantiopsida): total-evidence dating analysis supports a Late Silurian-Early Devonian origin and post-Mesozoic morphological stasis.

Divergence times based on molecular clock analyses often differ from those derived from total-evidence dating (TED) approaches. For bryophytes, fossils have been excluded from previous assessments of divergence times, and thus, their utility in dating analyses remains unexplored. Here, we conduct the first TED analyses of the complex thalloid liverworts (Marchantiopsida) that include fossils and evaluate macroevolutionary trends in morphological 'diversity' (disparity) and rates. Phylogenetic analyses were performed on a combined dataset of 130 discrete characters and 11 molecular markers (sampled from nuclear, plastid and mitochondrial genomes). Taxon sampling spanned 56 extant species - representing all the orders within Marchantiophyta and extant genera within Marchantiales - and eight fossil taxa. Total-evidence dating analyses support the radiation of Marchantiopsida during Late Silurian-Early Devonian (or Middle Ordovician when the outgroup is excluded) and that of Ricciaceae in the Middle Jurassic. Morphological change rate was high early in the history of the group, but it barely increased after Late Cretaceous. Disparity-through-time analyses support a fast increase in diversity until the Middle Triassic (c. 250 Ma), after which phenotypic evolution slows down considerably. Incorporating fossils in analyses challenges previous assumptions on the affinities of extinct taxa and indicates that complex thalloid liverworts radiated c. 125 Ma earlier than previously inferred.

From the morphospace to the soundscape: Exploring the diversity and functional morphology of the fish inner ear, with a focus on elasmobranchsa).

Fishes, including elasmobranchs (sharks, rays, and skates), present an astonishing diversity in inner ear morphologies; however, the functional significance of these variations and how they confer auditory capacity is yet to be resolved. The relationship between inner ear structure and hearing performance is unclear, partly because most of the morphological and biomechanical mechanisms that underlie the hearing functions are complex and poorly known. Here, we present advanced opportunities to document discontinuities in the macroevolutionary trends of a complex biological form, like the inner ear, and test hypotheses regarding what factors may be driving morphological diversity. Three-dimensional (3D) bioimaging, geometric morphometrics, and finite element analysis are methods that can be combined to interrogate the structure-to-function links in elasmobranch fish inner ears. In addition, open-source 3D morphology datasets, advances in phylogenetic comparative methods, and methods for the analysis of highly multidimensional shape data have leveraged these opportunities. Questions that can be explored with this toolkit are identified, the different methods are justified, and remaining challenges are highlighted as avenues for future work.

How Aestivation Evolved in Turtles: A Macroevolutionary and Morphological Approach

AbstractAestivation (summer dormancy) is a long-term multiday torpor in response to hot and dry periods. It has been detected in many species of terrestrial and aquatic turtles; however, several ecological and evolutionary aspects of chelonian aestivation remain to be evaluated and understood. We conducted a comparative exploration of macroevolutionary trends in turtle aestivation and tested the potential correlation of shell morphology with the aestivation duration. We compiled a dataset of aestivation status, aestivation times, and measurements of shell morphology of 225 turtle species. We reconstructed ancestral states along a time-calibrated phylogeny and tested different evolutionary models on the presence/absence of the aestivation trait. We also performed phylogenetic comparative analysis to explore several shell morphological traits likely associated with the duration time in aestivation behavior. We found evidence of aestivation in 44% of the evaluated turtle species. Aestivation times were longer in Chelidae, Pelomedusidae, Geoemydidae, and Kinosternidae, and the shortest times in Emydidae and Testudinidae. Aestivation behavior is a derived trait evolved independently and several times in the pleurodires and cryptodires turtle groups. We found some evolutionary trends in different turtle families, Pelomedusidae and Kinosternidae showed considerable increases in the presence of the aestivation trait, while families such as Podocnemididae, Trionychidae and Chelydridae showed important deductions for the same trait. Our results for the association between shell morphology and aestivation duration in turtles were contrasting among families. Overall, it was the contribution of several and different morphological traits that allow a positive and significant association with the aestivation times.

Deep resilience: An evolutionary perspective on calcification in an age of ocean acidification.

The success of today's calcifying organisms in tomorrow's oceans depends, in part, on the resilience of their skeletons to ocean acidification. To the extent this statement is true there is reason to have hope. Many marine calcifiers demonstrate resilience when exposed to environments that mimic near-term ocean acidification. The fossil record similarly suggests that resilience in skeletons has increased dramatically over geologic time. This "deep resilience" is seen in the long-term stability of skeletal chemistry, as well as a decreasing correlation between skeletal mineralogy and extinction risk over time. Such resilience over geologic timescales is often attributed to genetic canalization-the hardening of genetic pathways due to the evolution of increasingly complex regulatory systems. But paradoxically, our current knowledge on biomineralization genetics suggests an opposing trend, where genes are co-opted and shuffled at an evolutionarily rapid pace. In this paper we consider two possible mechanisms driving deep resilience in skeletons that fall outside of genetic canalization: microbial co-regulation and macroevolutionary trends in skeleton structure. The mechanisms driving deep resilience should be considered when creating risk assessments for marine organisms facing ocean acidification and provide a wealth of research avenues to explore.

The early diversification of ray-finned fishes (Actinopterygii): hypotheses, challenges and future prospects.

Actinopterygii makes up half of living vertebrate diversity, and study of fossil members during their Palaeozoic rise to dominance has a long history of descriptive work. Although research interest into Palaeozoic actinopterygians has increased in recent years, broader patterns of diversity and diversity dynamics remain critically understudied. Past studies have investigated macroevolutionary trends in Palaeozoic actinopterygians in a piecemeal fashion, variably using existing compendia of vertebrates or literature-based searches. Here, we present a comprehensive occurrence-based dataset of actinopterygians spanning the whole of the Palaeozoic. We use this to produce the first through-Palaeozoic trends in genus and species counts for Actinopterygii. Diversity through time generally tracks metrics for sampling, while major taxonomic problems pervading the Palaeozoic actinopterygian record obscure diversity trends. Many described species are concentrated in several particularly problematic 'waste-basket' genera, hiding considerable morphological and taxonomic diversity. This taxonomic confusion also feeds into a limited understanding of phylogenetic relationships. A heavy sampling bias towards Europe and North America exists in both occurrence databases and available phylogenetic matrices, with other regions underrepresented despite yielding important data. Scrutiny of the extent to which spatial biases influence the actinopterygian record is lacking, as is research on other forms of bias. Low richness in some time periods may be linked to geological biases, while the effects of taphonomic biases on Palaeozoic actinopterygians have not yet been investigated. Efforts are already underway both to redescribe poorly defined taxa and to describe taxa from underrepresented regions, helping to address taxonomic issues and accuracy of occurrence data. New methods of sampling standardisation utilising up-to-date occurrence databases will be critical in teasing apart biological changes in diversity and those resulting from bias. Lastly, continued phylogenetic work will enable the use of phylogenetic comparative methods to elucidate the origins of actinopterygian biogeography and subsequent patterns of radiation throughout their rise to dominate aquatic faunas.

Convergence, divergence, and macroevolutionary constraint as revealed by anatomical network analysis of the squamate skull, with an emphasis on snakes

Traditionally considered the earliest-diverging group of snakes, scolecophidians are central to major evolutionary paradigms regarding squamate feeding mechanisms and the ecological origins of snakes. However, quantitative analyses of these phenomena remain scarce. Herein, we therefore assess skull modularity in squamates via anatomical network analysis, focusing on the interplay between ‘microstomy’ (small-gaped feeding), fossoriality, and miniaturization in scolecophidians. Our analyses reveal distinctive patterns of jaw connectivity across purported ‘microstomatans’, thus supporting a more complex scenario of jaw evolution than traditionally portrayed. We also find that fossoriality and miniaturization each define a similar region of topospace (i.e., connectivity-based morphospace), with their combined influence imposing further evolutionary constraint on skull architecture. These results ultimately indicate convergence among scolecophidians, refuting widespread perspectives of these snakes as fundamentally plesiomorphic and morphologically homogeneous. This network-based examination of skull modularity—the first of its kind for snakes, and one of the first to analyze squamates—thus provides key insights into macroevolutionary trends among squamates, with particular implications for snake origins and evolution.

A database of the morphology, ecology and literature of the world's limb‐reduced skinks

AbstractAimLimb‐reduced squamates are a convenient model system to investigate macroevolutionary trends in morphology. Here, we provide morphological, ecological and literature data on all known species of limb‐reduced skinks (Scincidae) and their relatives, representing one of the most diverse and widely distributed groups of limb‐reduced squamates.LocationGlobal.TaxonSkinks (Reptilia, Squamata: Scincidae). Limb‐reduced forms.MethodsMorphological data were sourced from the primary literature, spanning a period of over 150 years. Linear body measurements were averaged across all values in the literature, preserving proportionality to body length. For digits and presacral vertebrae, we used maximum recorded counts. Ecological and biogeographical data were sourced from habitat assessments in the primary literature, online databases and field guides. Literature data were sorted according to type of study. To exemplify the applicability of the database, we used Markov‐chain ordered models to estimate the evolutionary frequency of limb reduction and loss in skinks.ResultsWe find evidence of limb reduction and loss in a total of 394 species worldwide, representing ~23% of all skink species, and ~30% of genera. The distribution of limb‐reduced and limbless forms differs from that of fully limbed forms, as they are present in all biogeographic realms with the almost complete exclusion of the Americas. We estimate that limb reduction evolved more than 50 times in skinks, and that loss of at least one limb pair evolved at least 24 times.Main conclusionsThe dataset captures a broad spectrum of morphological and ecological variation in a large, globally distributed taxonomic group. It establishes a widely applicable definition of limb reduction based on limb proportions as a reference for future studies. Such an extensive collection of morphological and ecological data can pave the way for investigations of dramatic morphological transitions and their ecological drivers at a global and local scale.

The chondrichthyan fossil record of the Florida Platform (Eocene–Pleistocene)

AbstractAs of September 2019, the Florida Museum of Natural History (FLMNH) had a curated collection of 117,449 chondrichthyan specimens from Florida, spanning the Eocene through the Pleistocene. Herein, I evaluate the completeness of the chondrichthyan fossil record from Florida based on the FLMNH collection, while analyzing patterns in taxonomic and ecomorphological diversity. At least 70 chondrichthyan taxa were recognized, representing 10 orders, 26 families, and 42 genera; of which, 20 taxa represent first occurrences from Florida. A sample of 107,698 specimens was organized into 12 time bins to analyze taxonomic and ecomorphological diversity, with an expectation that diversity patterns would correspond with global climate events (e.g., the Eocene–Oligocene transition and the middle Miocene climatic optimum). However, diversity patterns were obscured by pervasive sampling bias, attributable to variable collection methods, research prioritizations, and regional lithologic controls. Sampling is particularly poor for smaller specimens and older geologic units (e.g., the Paleogene). Despite incomplete sampling of the Florida chondrichthyan fossil record, there was an apparent turnover along the Atlantic and Gulf Coastal Plains from a lamniform- to carcharhiniform-dominated chondrichthyan fauna that occurred during the Eocene. This turnover corresponded with the extinction of many lamniform taxa with grasping-dominated dentition types (e.g., Brachycarcharias, Jaekelotodus, and Macrorhizodus). Selachian taxa that survived the late Eocene extinctions were predominantly represented by cutting-dominant dentition types. As cutting aids in the dismemberment of prey, this may reflect a macroevolutionary trend toward active predation and scavenging on larger prey, such as marine mammals, teleost fish, and other sharks.

A coherent biogeographical framework for Old World Neogene and Pleistocene mammals

AbstractIn order to understand mammalian evolution and compute a wide range of biodiversity indices, we commonly use the ‘bioregion’, a spatial division adapted to ecological and evolutionary constraints. While commonly conducted by neontologists, the establishment of bioregions in palaeontology is generally a secondary analysis, shaped on subjective time scales and areas specific to the investigated questions and groups. This heterogeneity, coupled with the scale‐dependency of biodiversity indices, prevents the clear identification of macroecological and macroevolutionary trends for large taxonomic groups like extinct mammals. Here we tackle this issue by providing a coherent framework for Neogene and Pleistocene mammals of the Old World following two steps: (1) a temporal scale adapted to mammalian evolutionary history (i.e. evolutionary fauna) is defined by poly‐cohort analysis; (2) bioregions are then computed for each evolutionary fauna by clustering, ordination and intermediate approaches at multiples spatial scales (i.e. continental to regional) for Eurasia and Africa. Additionally, providing a coherent framework for a wide range of mammalian datasets, our results show: (1) the synchronous emergence and fall of five mammalian evolutionary faunas identified at chronological scales varying from the epoch to the geological stage; (2) a transition from a longitudinal to a latitudinal biogeographical structuring between the Miocene and Pliocene, especially in Europe; (3) the long‐term affinity of southern Asian with African faunas, in sharp contrast with the modern Palaearctic bioregion extension; and (4) the establishment of a vast Mediterranean bioregion from fragmented areas in the Late Miocene to its full extent in the Pleistocene.

The macroevolutionary and developmental evolution of the turtle carapacial scutes

The scutes of the carapace of extant turtles exhibit common elements in a narrow range of topographical arrangements. The typical arrangement has remained constant since its origin in the clade Mesochelydia (Early Jurassic), after a period of apparent greater diversity in the Triassic. This contribution is a review of the development and evolutionary history of the scute patterns of the carapace, seen through the lens of recent developmental models. This yields insights on pattern variations in the fossil record. We reinterpret the “supracaudal” scute and propose thatProganochelyshad five vertebral scutes. We discuss the relationship between supramarginal scutes and Turing processes, and we show how a simple change during embryogenesis could account for origin of the configuration of the caudal region of the carapace in mesochelydians. We also discuss the nature of the decrease in number of scutes over the course of evolution, and whether macroevolutionary trends can be discerned. We argue that turtles with complete loss of scutes (e.g., softshells) follow clade-specific macroevolutionary regimes, which are distinct from the majority of other turtles. Finally, we draw a parallel between the variation of scute patterns on the carapace of turtles and the scale patterns in the pileus region (roof of the head) of squamates. The size and numbers of scales in the pileus region can evolve over a wide range, but we recognized tentative evidence of convergence towards a typical configuration when the scales become larger and fewer. Thus, typical patterns could be a more general property of similar systems of integumentary appendages.

Review of morphology, development, and evolution of the notarium in birds.

The notarium is a rigid bony structure, which resulted from the fusion of thoracic vertebrae of some pterosaurs and birds. It is high variable, ranging from two to six fused thoracic vertebrae. In this study, we reviewed and analyzed approximately 270 specimens of neornithine birds (representing 80% of the living orders) and some fossils in order to identify the number of fused vertebrae, degree and sites of vertebral fusion, occurrence of sutures, and other structures of potential phylogenetic and functional significance. These data were analyzed using a recent time-calibrated molecular phylogenetic tree and principal component analyses analysis evaluating the relationship with long bones in order to reconstruct macroevolutionary trends related to the evolution of the notarium. The occurrence of this structure shows a mosaic distribution over neornithine phylogeny, originating several times independently, especially during the Paleogene, in predominantly ground-dwelling forms. The notarium of these groups is characterized by: neural spines fused into single structure, intervertebral openings small to absent, large ventral keels forming ventral plates, and fused transverse processes. Derived neornithines, such as aquatic forms and long-legged birds, have a tendency to display a decreased degree of fusion between the vertebrae, which may indicate a reduction or disappearance of the notarium.

Macroevolutionary trends in theropod dinosaur feeding mechanics

Theropod dinosaurs underwent some of the most remarkable dietary changes in vertebrate evolutionary history, shifting from ancestral carnivory1-3 to hypercarnivory4,5 and omnivory/herbivory,6-9 with some taxa eventually reverting to carnivory.10-12 The mandible is an important tool for food acquisition in vertebrates and reflects adaptations to feeding modes and diets.13,14 The morphofunctional modifications accompanying the dietary changes in theropod dinosaurs are not well understood because most of the previous studies focused solely on the cranium and/or were phylogenetically limited in scope,12,15-21 while studies that include multiple clades are usually based on linear measurements and/or discrete osteological characters.8,22 Given the potential relationship between macroevolutionary change and ontogenetic pattern,23 we explore whether functional morphological patterns observed in theropod mandibular evolution show similarities to the ontogenetic trajectory. Here, we use finite element analysis to study the mandibles of non-avialan coelurosaurian theropods and demonstrate how feeding mechanics vary between dietary groups and major clades. We reveal an overall reduction in feeding-induced stresses along all theropod lineages through time. This is facilitated by a post-dentary expansion and the development of a downturned dentary in herbivores and an upturned dentary in carnivores likely via the "curved bone effect." We also observed the same reduction in feeding-induced stress in an ontogenetic series of jaws of the tyrannosaurids Tarbosaurus and Tyrannosaurus, which is best attributed to bone functional adaptation. This suggests that this common tendency for structural strengthening of the theropod mandible through time, irrespective of diet, is linked to "functional peramorphosis" of bone functional adaptations acquired during ontogeny.

General patterns of sexual dimorphism in graylings (Thymallus), with a comparison to other salmonid species

Among fishes, salmonids (family Salmonidae) have attracted a great deal of research attention focused on sexual dimorphism and associated selective forces. Most of this research has been directed toward anadromous and mostly semelparous salmon and trout (Oncorhynchus, Salmo), and comparatively little is known about intersexual variability in strictly iteroparous freshwater salmonids. We examined a comprehensive data set of 28 linear morphometric characters in 11 of 15 currently recognised species of grayling (Thymallinae, Thymallus), a genus consisting of iteroparous species only, to identify general patterns of intersexual morphological variability. Overall, we found that all grayling species show common sex-specific traits particularly relating to size dimensions of the dorsal, anal, pelvic and pectoral fins. Although the magnitude of sexual dimorphism differed among species, there was no significant phylogenetic signal associated with these differences across the genus. These results are discussed in terms of the assumed selection pressures driving sexual dimorphism in graylings and are compared to existing knowledge in Salmonidae as a whole where similarities and differences with both Salmoninae and Coregoninae exist. The present study provides the first detailed genus-wide comparison of sexually dimorphic phenotypic characters in graylings, and highlights the need for more large-scale comparative studies in multiple salmonid species to better understand general macroevolutionary trends among this important group of freshwater fishes.

No evidence for parallel evolution of cursorial limb adaptations among Neogene South American native ungulates (SANUs).

During the Neogene, many North American ungulates evolved longer limbs. Presumably, this allowed them to move more efficiently or quickly in open habitats, which became more common during this interval. Evidence suggests that open habitats appeared even earlier in South America, but no study to date has investigated whether the ungulate-like mammals of South America (South American native ungulates or SANUs) evolved similar limb adaptations. We analyzed limb elongation in the two predominant SANU groups, notoungulates and litopterns, by compiling genus-level occurrences from the late Oligocene to the Pleistocene and calculating metatarsal/femur ratio (Mt:F). None of the groups or subgroups we analyzed show a pronounced increase in Mt:F across this interval, with the possible exception of proterotheriid litopterns. Proterotheriids are thought to have inhabited forested environments rather than open ones, which raises questions about the selective forces responsible for limb elongation in ungulates. Conversely, notoungulates, which are traditionally thought to have lived in open habitats, show no strong trend of increasing Mt:F across this interval. Our study suggests that the macroevolutionary trend of limb elongation in ungulate-like mammals is not universal and is highly influenced by the evolutionary affinities of the groups being analyzed.