Phylogeny, neoteny and growth of the cranial base in hominoids

This study tests the hypothesis that there is a general pattern in the growth of the cranial base of Homo sapiens that is 'essentially neotenous' [Gould, 1977]. Juvenile and adult crania of Homo sapiens, Gorilla gorilla, Pan troglodytes and Pongo pygmaeus were studied and the cross-sectional growth curves for 10 measurements made on the cranial base (as viewed in norma basilaris) were compared. The results of this study suggest that relatively simple modifications to the timing or pattern of growth are insufficient to explain the observed morphological differences between the cranial base of modern Homo sapiens and the great apes.     

Phylogeny, integration and expression of sigma virus-like genes in Drosophila

The recent and surprising discovery of widespread NIRVs (non-retroviral integrated RNA viruses) has highlighted the importance of genomic interactions between non-retroviral RNA viruses and their eukaryotic hosts. Among the viruses with integrated representatives are the rhabdoviruses, a family of negative sense single-stranded RNA viruses. We identify sigma virus-like NIRVs of Drosophila spp. that represent unique cases where NIRVs are closely related to exogenous RNA viruses in a model host organism. We have used a combination of bioinformatics and laboratory methods to explore the evolution and expression of sigma virus-like NIRVs in Drosophila. Recent integrations in Drosophila provide a promising experimental system to study functionality of NIRVs. Moreover, the genomic architecture of recent NIRVs provides an unusual evolutionary window on the integration mechanism. For example, we found that a sigma virus-like polymerase associated protein (P) gene appears to have been integrated by template switching of the blastopia-like LTR retrotransposon. The sigma virus P-like NIRV is present in multiple retroelement fused open reading frames on the X and 3R chromosomes of Drosophila yakuba - the X-linked copy is transcribed to produce an RNA product in adult flies. We present the first account of sigma virus-like NIRVs and the first example of NIRV expression in a model animal system, and therefore provide a platform for further study of the possible functions of NIRVs in animal hosts.     

Post-weaning cranial ontogeny in two bandicoots (Mammalia,Peramelomorphia, Peramelidae) and comparison with carnivorous marsupials

The ontogeny of the skull has been studied in several marsupial groups such as didelphids, microbiotheriids, and dasyurids. Here, we describe and compare the post-weaning ontogeny of the skull in two species of bandicoots, Echymipera kalubu (Echymiperinae) and Isoodon macrourus (Peramelinae), analyzing specific allometric trends in both groups, describing common (and specific) patterns, and discussing them on functional and phylogenetic grounds. Growth patterns were analyzed both qualitatively and quantitatively, including bivariate and multivariate analyses of allometry. We also evaluated character transformation and phylogenetic signals of the allometric patterns in several groups of marsupials and some placentals. We identified morphological changes between juvenile and adult stages in both species of peramelids, many related to the development of the trophic apparatus. Notable differences were detected in the patterns of growth, suggesting divergences in ontogenetic trajectories between both species. Both bivariate and multivariate methods indicate that positive allometries in E. kalubu apply to longitudinal dimensions, whereas in I. macrourus, positive allometries are restricted to vertical dimensions of the skull. The comparison of the allometric trends of two bandicoots with previously studied taxa reveals that although peramelids exhibit a particularly short gestation period and divergent morphology compared to other marsupials, their pattern does not show any particular trend. Some allometric trends seem to be highly conserved among the species studied, showing weak phylogenetic signal. Marsupials in general do not show particular patterns of post-weaning skull growth compared with placentals.     

Timing of cranial suture closure in placental mammals: Phylogenetic patterns,intraspecific variation,and comparison with marsupials

Used as markers of postnatal growth closure sequences of 22 ectocranial sutures and synchondroses were recorded in a sample of 1161 skulls belonging to 38 species from all major placental clades: Afrotheria, Xenarthra, Laurasiatheria and Euarchontoglires (Boreoeutheria). The maximum closure level, which is not significantly correlated to body mass, is higher in Afrotheria and Xenarthra than in Boreoeutheria. Only the basioccipito‐basisphenoid and the basioccipito‐exoccipital synchondroses close in all species sampled, the supraoccipito‐exoccipital and the inter‐parietal sutures do in most species. Parsimov retrieved more heterochronic shifts for Afrotheria and Xenarthra than for Boreoeutheria. The amount of intraspecific variation differs among the species sampled being high among xenarthran species and low among afrotherians. Specimens (162) representing 12 marsupial genera were also analysed. Placentals exhibit a larger number of suture closures than marsupials and in both groups the sutures at the base of the skull are the first to fuse starting with the basioccipito‐exoccipital. J. Morphol. 275:125–140, 2014. © 2013 Wiley Periodicals, Inc.     

Phylogeny and evolution of body mass in didelphid marsupials (Marsupialia: Didelphimorphia: Didelphidae)

Most extant New World marsupials belong in the Didelphidae, which comprises ca. 110 currently recognized species of opossums. Didelphids are small mammals with their mean body mass, at species level, ranging from ca. 7 g to 2.2 kg. The largest species belong in a single clade, while substantial variation remains scattered across the remaining groups. We seek out to explore the details of this mass variation in an evolutionary framework. To this end, we first reconstructed the phylogeny of didelphids based on an extensive, although fragmentary sample of sequences from ten genes. We recovered a fully resolved, highly robust phylogeny that tested and confirmed most previously reported groupings, providing a simultaneous depiction of phylogenetic relationships for 81 % of currently recognized species and all relevant supra-specific clades. As much as 69 % of total body mass variation in didelphids was explained by this phylogenetic hypothesis. Mapped on it, mass variation evolved as much as 6.8 kg of total changes, starting from a reconstructed ancestral body mass range of 22–33 g. No single, family-wide pattern was evident; in fact, the dominant pattern for mass variation was that of increases in body mass along a few successive branches, or phyletic giantism, followed by apomorphic nanism, i.e., decreases localized in single terminal branches. Phyletic trends indicated the persistence of gradual, directional changes along considerable spans of geological time and show that substantial variation of interest resides in this and perhaps most groups of small mammals.     

Morphological evolution through integration: a quantitative study of cranial integration in Homo, Pan, Gorilla and Pongo

Morphological integration refers to coordinated variation among traits that are closely related in development and/or function. Patterns of integration can offer important insight into the structural relationship between phenotypic units, providing a framework to address questions about phenotypic evolvability and constraints. Integrative features of the primate cranium have recently become a popular subject of study. However, an important question that still remains under-investigated is: what is the pattern of cranial shape integration among closely related hominoids? To address this question, we conducted a Procrustes-based geometric morphometrics study to quantify and analyze shape covariation patterns between different cranial regions in Homo, Pan, Gorilla and Pongo. A total of fifty-six 3D landmarks were collected on 407 adult individuals. We then sub-divided the landmarks corresponding to cranial units as outlined in the ‘functional matrix hypothesis.’ Sub-dividing the cranium in this manner allowed us to explore patterns of covariation between the face, basicranium and cranial vault, using the two-block partial least squares approach. Our results suggest that integrated shape changes in the hominoid cranium are complex, but that the overall pattern of integration is similar among human and non-human apes. Thus, despite having very distinct morphologies the way in which the face, basicranium and cranial vault covary is shared among these taxa. These results imply that the pattern of cranial integration among hominoids is conserved.     

Comparative finite element analysis of the cranial performance of four herbivorous marsupials

Marsupial herbivores exhibit a wide variety of skull shapes and sizes to exploit different ecological niches. Several studies on teeth, dentaries, and jaw adductor muscles indicate that marsupial herbivores exhibit different specializations for grazing and browsing. No studies, however, have examined the skulls of marsupial herbivores to determine the relationship between stress and strain, and the evolution of skull shape. The relationship between skull morphology, biomechanical performance, and diet was tested by applying the finite element method to the skulls of four marsupial herbivores: the common wombat (Vombatus ursinus), koala (Phascolarctos cinereus), swamp wallaby (Wallabia bicolor), and red kangaroo (Macropus rufus). It was hypothesized that grazers, requiring stronger skulls to process tougher food, would have higher biomechanical performance than browsers. This was true when comparing the koala and wallaby (browsers) to the wombat (a grazer). The cranial model of the wombat resulted in low stress and high mechanical efficiency in relation to a robust skull capable of generating high bite forces. However, the kangaroo, also a grazer, has evolved a very different strategy to process tough food. The cranium is much more gracile and has higher stress and lower mechanical efficiency, but they adopt a different method of processing food by having a curved tooth row to concentrate force in a smaller area and molar progression to remove worn teeth from the tooth row. Therefore, the position of the bite is crucial for the structural performance of the kangaroo skull, while it is not for the wombat which process food along the entire tooth row. In accordance with previous studies, the results from this study show the mammalian skull is optimized to resist forces generated during feeding. However, other factors, including the lifestyle of the animal and its environment, also affect selection for skull morphology to meet multiple functional demands. J. Morphol. 276:1230–1243, 2015. © 2015 Wiley Periodicals, Inc.     

The evolution of cranial design, diet, and feeding mechanisms in batoid fishes

The batoid fishes (electric rays, sawfishes, skates, guitarfishes,and stingrays) are a trophically and morphologically diverseclade in which the observed range of diets is a product of afeeding mechanism with few parts and therefore a limited numberof functional interactions. This system allows an intriguingcomparison to the complex network of associations in the feedingapparatus of bony fishes and an anatomically simple frameworkfor investigations of the mechanisms underlying the evolutionof functional and phenotypic diversity. We quantified morphologyfrom reconstructed CT scans of 40 batoid species, representingmore than half of the extant genera. We used pairwise comparisonsto evaluate the extent of coevolution among components of thefeeding apparatus and among morphologies and diets. These relationshipswere then used to predict diets in poorly studied taxa and ina reconstruction of the batoid ancestor. Although functionallythere are fewer examples of convergence in the batoid feedingmechanism than in bony fishes, our data show multiple evolutionsof similar dietary compositions underlain by a broad morphologicaldiversity. Elements of the feeding apparatus evolved independentlyof one another, suggesting that decoupling components of thehead skeleton created separate but interacting evolutionarymodules that allowed trophic diversification. Our data implythat food habits exhibit strong independent and convergent evolutionand that suites of morphologies are associated with certaindiets; however, lack of behavioral data for this clade, andone example of divergent diets underlain by convergent morphology,caution against the assumption of simplistic relationships betweenform and function. We therefore urge future work to ground truthour study by testing the functional, dietary and evolutionaryhypotheses suggested by our data.     

A comparative analysis of the evolutionary relationship between diet and enzyme targeting in bats, marsupials and other mammals

The subcellular distribution of the enzyme alanine:glyoxylate aminotransferase (AGT) in the livers of different mammals appears to be related to their natural diets. Thus, AGT tends to be mitochondrial in carnivores, peroxisomal in herbivores, and both mitochondrial and peroxisomal in omnivores. To what extent this relationship is an incidental consequence of phylogenetic structure or an evolutionarily meaningful adaptive response to changes in dietary selection pressure is unknown. In order to distinguish between these two possibilities, we have determined the subcellular distribution of AGT in the livers of 22 new mammalian species, including members of three orders not studied before. In addition, we have analysed the statistical relationship between AGT distribution and diet in all 77 mammalian species, from 12 different orders, for which the distribution is currently known. Our analysis shows that there is a highly significant correlation between AGT distribution and diet, independent of phylogeny. This finding is compatible with the suggestion that the variable intracellular targeting of AGT is an adaptive response to episodic changes in dietary selection pressure. To our knowledge, this is the first example of such a response being manifested at the molecular and cellular levels across the breadth of Mammalia.     

Correlations between lizard cranial shape and diet: a quantitative, phylogenetically informed analysis

Although the relationship between dietary and phenotypic specialization has been well documented for many vertebrate groups, it has been stated that few such general trends can be established for lizards. This is often thought to be due to the lack of dietary specialization in many lizards. For example, many species that are reported to be insectivorous may also consume a variety of plant materials, and the reverse is often true as well. In this study, we investigate whether a correlation exists between general cranial form and dietary niche in lizards. Additionally, we test previously proposed hypotheses suggesting that herbivorous lizards should be larger bodied than lizards with other diets. Our data indicate that lizards specializing in food items imposing different mechanical demands on the feeding system show clear patterns of morphological specialization in their cranial morphology. True herbivores (diet of fibrous and tough foliage) are clearly distinguished from omnivorous and carnivorous lizards by having taller skulls and shorter snouts, likely related to the need for high bite forces. This allows herbivores to mechanically reduce relatively less digestible foliage. Carnivores have relatively longer snouts and retroarticular processes, which may result in more efficient capture and processing of elusive prey. When analysed in an explicit phylogenetic context, only snout length and skull mass remained significantly different between dietary groups. The small number of differences in the phylogenetic analyses is likely the result of shared evolutionary history and the relative paucity of independent origins of herbivory and omnivory in our sample. Analyses of the relationship between diet and body size show that on average herbivores have a larger body size than carnivores, with omnivores intermediate between the two other dietary groups. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 433–466.     

Birth in marsupials

Birth is an event that allows the relatively immature marsupial to move from the internal environment of the uterus to the external environment of the pouch. The newborn marsupial passes down from the uterus to the urogenital sinus and then makes its way to the pouch and attaches to the teat at a very early stage of development. From the studies available, there appear to be three methods used by the newborn to move from the uterus to the pouch. In marsupials with a forward pouch such as the red kangaroo, tammar wallaby and the brushtail possum, the mother positions her urogenital sinus below the pouch and the newborn climb upward towards the pouch. The young climb with a swimming motion, moving the head from side-to-side and use the forearms in alternate strokes. In the bandicoot with a backward facing pouch, the mother positions the urogenital sinus above the pouch and the young slither down into the pouch. The young do not have a definite crawl, as seen with the macropodids and possum. The third method of birth has been observed in the marsupials without a definite pouch that have a mammary region that develops as the young grow in size. This type of pouch is observed in the dasyurids. The mother was noted to stand on four legs with her hips raised so that the urogenital sinus was above the pouch and the newborn young crawled downwards from the sinus to the pouch. In all species, birth was completed in 2–4 min.     

Comparison of cranial form and function in association with diet in natricine snakes

The skull of squamates has many functions, with food acquisition and ingestion being paramount. Snakes vary interspecifically in the frequency, size, and types of prey that are consumed. Natural selection should favor phenotypes that minimize the costs of energy acquisition; therefore, trophic morphology should reflect a snake's primary prey type to enhance some aspect of feeding performance. I measured 19 cranial variables for six natricine species that vary in the frequency with which they consume frogs and fish. Both conventional and phylogenetically corrected analyses indicated that fish‐eating snakes have relatively longer upper and lower jaw elements than frog‐eating snakes, which tended to have broader skull components. I also compared the ratio of the in‐lever to the out‐lever lengths of the jaw‐closing mechanism [jaw mechanical advantage (MA)] among species. Fish‐eating snakes had significantly lower MAs in the jaws than did the frog‐eating snakes. This result suggests that piscivores have faster closing jaws and that the jaws of frog‐eating snakes have higher closing forces. Cranial morphology and the functional demands of prey capture and ingestion appear to be associated with primary prey type in natricine snakes. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

Food habits, energetics, and the reproduction of marsupials

Basal rate of metabolism in marsupials and in eutherian mammals is principally correlated with body mass, food habits and activity. Feeding on fruit, the leaves of woody plants, or invertebrates is associated with low basal rates, especially at large masses, in both groups of mammals. These foods lead to low basal rates because they are seasonally unavailable, are indigestible, or need to be detoxified. The depression in basal rate associated with frugivory and folivory is increased when coupled with sedentary, arboreal habits in both marsupials and eutherians. In contrast, eutherians that feed on vertebrates or herbs generally have high basal rates, while marsupials that eat these foods do not have high basal rates. These foods permit high basal rates, which are exploited by eutherians because high basal rates in these mammals lead to high rates of reproduction. Marsupials have, at best, a limited correlation of reproduction with rate of metabolism, so that feeding on vertebrates or herbs does not lead to high basal rates in these mammals. This difference between marsupials and eutherians in the coupling of reproduction to energetics has at least two ecological consequences. 1) Marsupials generally do not tolerate cold-temperate environments because they do not accelerate growth and development to complete reproduction within a short spring and summer. 2) Marsupials coexist with ecologically similar eutherians as long as marsupials have food habits that are correlated with low rates of metabolism in eutherians (i.e. they feed on fruit, the leaves of woody plants, or invertebrates), but they tend to be displaced by eutherians when marsupials have food habits that are associated with high rates of metabolism in eutherians (i.e. when they feed on vertebrates and, probably, herbs).     

Size-related changes in cranial morphology affect diet in the catfish Clariallabes longicauda

Within the catfish family Clariidae, species exist with different degrees of jaw adductor hypertrophy. This jaw adductor hypertrophy has been related to bite performance, in turn suggesting a link to dietary specialization. Thus, an increase in the degree of hypertrophy will likely be reflected in an increase in the amount of hard prey in the diet. In the present study, we examine the ontogenetic scaling of cranial structure and diet in a species of catfish with a moderate degree of jaw adductor hypertrophy, Clariallabes longicauda . Additionally, we investigate whether the observed changes in the morphology of the feeding system during growth are linked to changes in diet. The fish examined demonstrate a strong positively allometric growth of the jaw adductors, of head height and of maximal head width, suggesting that larger fish can feed on larger and harder prey. Dietary data confirm these hypotheses and reveal an increase in maximal prey size consumed, the proportion of large prey in the diet, and average prey hardness during ontogeny. Moreover, the observed changes in the proportion of large prey consumed and prey hardness are correlated with an increase in lower jaw width and maximal head width, respectively. An increase in the amount of evasive prey in the diet with fish size is correlated with an increase in hyoid length. In summary, not only size dependent, but also size-independent variation of the feeding system was associated with ontogenetic changes in diet in C. longicauda .  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 323–334.     

Integration,heterochrony, and adaptation in pedal digits of syndactylous marsupials

Background  

Marsupial syndactyly is a curious morphology of the foot found in all species of diprotodontian and peramelemorph marsupials. It is traditionally defined as a condition in which digits II and III of the foot are bound by skin and are reduced. Past treatments of marsupial syndactyly have not considered the implications of this unique morphology for broader issues of digit development and evolution, and the ongoing debate regarding its phylogenetic meaning lacks a broad empirical basis. This study undertakes the first interdisciplinary characterisation of syndactyly, using variance/covariance matrix comparisons of morphometric measurements, locomotor indices, ossification sequences, and re-assessment of the largely anecdotal data on the phylogenetic distribution of tarsal/metatarsal articulations and "incipient syndactyly".