Ecomorphological correlates of craniodental variation in bears and paleobiological implications for extinct taxa: an approach based on geometric morphometrics

Relative warp analyses of landmarks describing cranial and mandibular shape are used for investigating patterns of morphological variation among extant bears (Mammalia, Carnivora, Ursidae) indicative of diet and feeding behavior. These patterns are used for deriving inferences about the autecology of two extinct species previously assumed to have had different dietary preferences, the North American giant, short-faced bear Arctodus simus and the Eurasian cave bear Ursus spelaeus . Results reveal a set of shared craniodental traits among the herbivorous bears, including short and vaulted skulls with well-developed zygomatic arches, lateralized orbits and small canines, concave jaws with a highly positioned condyle, large moment arms for the temporalis and masseter muscles, and long cheek teeth. In contrast, those bears that consume animal resources have long skulls with small zygomatic arches, frontalized orbits and well-developed canines, and long jaws with a deep mandibular symphysis, low muscle leverages, a condyle situated at the level of the tooth row and reduced cheek teeth. The craniodental morphology of omnivorous bears is intermediate between those of faunivores and herbivores. This is also the case of the short-faced bear and the cave bear, which suggests that previous reconstructions of the feeding ecology of these extinct species (highly carnivorous for A. simus and herbivorous for U. spelaeus ) should be revised.     

A three-dimensional analysis of tooth-root morphology in living bears and implications for feeding behaviour in the extinct cave bear

ABSTRACT

The morphology of both crowns and tooth-roots reflects dietary specialisation in mammalian carnivores. In this article, we analyse the tooth-root morphology of maxillary teeth from CT scans of living bears (Ursus arctos, Ursus americanus, Ursus maritimus, Ursus thibetanus, Melursus ursinus, Helarctos malayanus, Tremarctos ornatus and Ailuropoda melanoleuca) in order to make inferences about the diet and feeding behaviour of the extinct cave bear (Ursus spelaeus sensu lato). Specifically, we investigate two major mitochondrial clades of extinct cave bears recognized by previous authors: Ursus ingressus and Ursus spelaeus (U. spelaeus spelaeus, U. spelaeus ladinicus, U. spelaeus eremus). Our results indicate a close association between tooth-root surface area and feeding behaviour in all living bear species. Tooth-root surface area values of cave bears suggest that they relied more on vegetative matter than living brown bears (Ursus arctos) but subtle differences between these species/subspecies could also indicate different feeding strategies among the members of cave bear complex.     

A new species of Carpolestes (Mammalia,Plesiadapoidea) from the late Paleocene of southern Wyoming: assessing changes in size and shape during the evolution of a key anatomical feature

A new and phylogenetically basal species of Carpolestes, the youngest and most derived genus of the plesiadapoid family Carpolestidae in North America, is described from a late Tiffanian (Ti-5) site in Sweetwater County, Wyoming, USA. Carpolestids differ from closely related plesiadapoid clades in having an enlarged, multicuspidate, blade-like P₄ that is partly convergent on that of multituberculates and other mammals showing plagiaulacoid dental adaptations. With some notable exceptions, the evolutionary history of North American carpolestids is characterized by the progressive development of larger and more elaborate P₄ blades through time. In particular, species of the monophyletic genus Carpolestes differ from species assigned to the earlier and apparently paraphyletic genus Carpodaptes in terms of both the size and shape of their P₄. A geometric morphometric analysis reveals that, with respect to P₄ shape, the closest approximation to the highly derived morphology of Carpolestes is made by Carpodaptes hobackensis, which is one of the smallest known species of Carpodaptes. In contrast, the largest known species of Carpodaptes, Carpodaptes jepseni, has a P₄ that falls within the metric range of variation for species of Carpolestes, yet Carpodaptes jepseni shows a uniquely derived P₄ shape that seems to exclude it from any special phylogenetic relationship with Carpolestes. A phylogenetic analysis based on dental characters reconstructs Carpodaptes hobackensis as the sister group of the Carpolestes clade. Shape seems to have been a more important factor than size during the final transformation of the blade-like P₄ of North American carpolestids.

http://zoobank.org/urn:lsid:zoobank.org:pub:212F9ECC-DA9A-44F8-BE7E-43F3EBAD636A     

Geographic pattern of cranial differentiation in the Asian Midday Jird Meriones meridianus (Rodentia: Muridae: Gerbillinae) and its taxonomic implications

The existence of cryptic species in the midday jird (Meriones meridianus) has been suggested in literature, although based on little empirical data to support this hypothesis. In this study, a two‐dimensional landmark‐based geometric morphometric approach was used to investigate whether patterns in intraspecific variation in skull shape and size exist, using 110 skull specimens from more than 20 different localities along the distribution range of M. meridianus. This is the first study of morphological differences on such a big sample size and geographical range, and it tries to find whether skull shape variation in this species is best described as being clinal or rather reflecting cryptic diversity. The latter seems to be the case, as a dimorphic skull phenotype was found, reflecting a geographic disparity between the Middle East and the Far East specimens both in shape and in size. Distinct cranial differences were found in the overall cranial size and, also at the level of the inflation of the bulla, the elongation of the nasal, the length of the teeth row and the incisive foramen, as well as the distance in between the latter two. It thus seems that M. meridianus from Middle East is morphologically distinct from that of the Far East. Furthermore, our results also demonstrate that clinal variation could explain variation within Middle East populations, whereas a more heterogenous pattern is found for those of the Far East. The hypothesis that the observed phenotypic variation may reflect cryptic species is discussed, with the recommendation for a thorough taxonomical revision of the genus in the region.     

Evolutionary divergence in Dolichopoda cave crickets: A comparison of single copy DNA hybridization data with allozymes and morphometric distances

In this paper we attempt to investigate relationships between the amount of genetic divergence in nuclear genes and the degree of morphological differentiation for different sets of characters in Dolichopoda cave crickets. Six populations representing five Dolichopoda species from Central and Southern Italy have been studied. The overall genetic divergence at nuclear genes was estimated both by single copy DNA-DNA hybridization and allozyme frequencies at 26 loci. Euclidean distances for two multivariate sets of morphometric variables: one describing body and appendage morphology, the other male epiphallus shape. Results showed a close agreement between the branching patterns of ΔTm values from DNA hybridization and Nei's allozyme distance values. On the other hand, patterns of morphological divergence revealed independent trends, although the branching pattern based on epiphallus morphology matched to some extent the phylogenies inferred from molecular data. The relative value of molecular and morphological characters as reliable phylogenetic tracers was evaluated in relation to their dependence on evolutionary factors. Implications of these findings on the calibration of molecular clocks are also discussed. The absolute rate of molecular change based on scDNA was estimated to be at least 0.98% divergence/my/lineage. This result is in agreement with calibrations attempted on other insects. Estimates of time of divergence based on allozymes (Nei's D) were highly consistent with the estimate from geological data.     

A new Middle Miocene tarsier from Thailand and the reconstruction of its orbital morphology using a geometric-morphometric method

Tarsius is an extant genus of primates endemic to the islands of Southeast Asia that is characterized by enormously enlarged orbits reflecting its nocturnal activity pattern. Tarsiers play a pivotal role in reconstructing primate phylogeny, because they appear to comprise, along with Anthropoidea, one of only two extant haplorhine clades. Their fossils are extremely rare. Here, we describe a new species of Tarsius from the Middle Miocene of Thailand. We reconstructed aspects of its orbital morphology using a geometric-morphometric method. The result shows that the new species of Tarsius had a very large orbit (falling within the range of variation of modern Tarsius) with a high degree of frontation and a low degree of convergence. Its relatively divergent lower premolar roots suggest a longer mesial tooth row and therefore a longer muzzle than in extant species. The new species documents a previous unknown Miocene group of Tarsius, indicating greater taxonomic diversity and morphological complexity during tarsier evolution. The current restriction of tarsiers to offshore islands in Southeast Asia appears to be a relatively recent phenomenon.     

Fighting the stigma caused by mental disorders: past perspectives,present activities,and future directions

People who live with mental illnesses are among the most stigmatized groups in society. In 1996, in recognition of the particularly harsh burden caused by the stigma associated with schizophrenia, the WPA initiated a global anti-stigma program, Open-the-Doors. In 2005, a WPA Section on Stigma and Mental Health was created, with a broader mandate to reduce stigma and discrimination caused by mental disabilities in general. In light of these impor-tant developments, and the growing public health interest in stigma reduction, this paper reflects on the past perspectives that have led us to our current position, reviews present activities and accomplishments, and identifies challenges that the Section members will face in their future efforts to reduce the stigma caused by mental disorders.     

Aerobic MTBE biodegradation: an examination of past studies, current challenges and future research directions

With the current practice of amending gasoline with up to 15% by volume MTBE, the contamination of groundwater by MTBE has become widespread. As a result, the bioremediation of MTBE-impacted aquifers has become an active area of research. A review of the current literature on the aerobic biodegradation of MTBE reveals that a number of cultures from diverse environments can either partially degrade or completely mineralize MTBE. MTBE is either utilized as a sole carbon and energy source or is degraded cometabolically by cultures grown on alkanes. Reported degradation rates range from 0.3 to 50 mg MTBE/g cells/h while growth rates (0.01–0.05 g MTBE/g cells/d) and cellular yields (0.1–0.2 g cells/g MTBE) are generally low. Studies on the mechanisms of MTBE degradation indicate that a monooxygenase enzyme cleaves the ether bond yielding tert-butyl alcohol (TBA) and formaldehyde as the dominant detectable intermediates. TBA is further degraded to 2-methyl-2-hydroxy-1-propanol, 2-hydroxyisobutyric acid, 2-propanol, acetone, hydroxyacteone and eventually, carbon dioxide. The majority of these intermediates are also common to mammalian MTBE metabolism. Laboratory studies on the degradation of MTBE in the presence of gasoline aromatics reveal that while degradation rates of other gasoline components are generally not inhibited by MTBE, MTBE degradation could be inhibited in the presence of more easily biodegradable compounds. Controlled field studies are clearly needed to elucidate MTBE degradation potential in co-contaminant plumes. Based on the reviewed studies, it is likely that a bioremediation strategy involving direct metabolism, cometabolism, bioaugmentation, or some combination thereof, could be applied as a feasible and cost-effective treatment method for MTBE contamination.     

Ontogeny and sex alter the effect of predation on body shape in a livebearing fish: sexual dimorphism, parallelism, and costs of reproduction

Predation can cause morphological divergence among populations, while ontogeny and sex often determine much of morphological diversity among individuals. We used geometric morphometrics to characterize body shape in the livebearing fish Brachyrhaphis rhabdophora to test for interactions between these three major shape-determining factors. We assessed shape variation between juveniles and adults of both sexes, and among adults for populations from high- and low-predation areas. Shape differed significantly between predation regimes for all juveniles regardless of sex. As males grew and matured into adults, ontogenetic shape trajectories were parallel, thus maintaining shape differences in adult males between predation environments. However, shape of adult females between predation environments followed a different pattern. As females grew and matured, ontogenetic shape trajectories converged so that shape differences were less pronounced between mature females in predator and nonpredator environments. Convergence in female body shape may indicate a trade-off between optimal shape for predator evasion versus shape required for the livebearing mode of reproduction.     

A new Stygonitocrella Petkovski (Copepoda: Harpacticoida) from a cave in Northern Mexico with comments on the taxonomy of the genus

The cave-dwelling harpacticoid copepod fauna of Mexico is still yet to be known. There are only a few records related to karstic caves, but most refer to epigean forms. The analysis of samples collected in a cave system in Northern Mexico yielded two female specimens of an unknown harpacticoid copepod species which is described herein. It was assigned to the genus Stygonitocrella Petkovski, 1976 mainly because of the presence of a single endopodal segment on legs 1–4; the new species, S. mexicana n. sp. differs from its congeners by a combination of characters related to the segmentation and armature of the swimming legs 1–4, the fifth leg of the female, and the strcture of the antennae. Based on the current limits of Stygonitocrella, two new subgenera are proposed to recognize different morphological patterns based on the presence or absence of an inner baseoendopodal lobe in the female fifth leg, a character not evaluated before. This lobe is absent in the new species and in other five species now contained in the subgenus Eustygonitocrella. A lobe is present in the new subgenus Fiersiella, also with six species. This is the second record of Stygonitocrella in North America, after S. (Fiersiella) sequoyahi Reid, Hunt & Stanley, 2003; it is the only North American cave-dwelling species of the genus. It is suggested that the influence of marine transgressions in Northern Mexico was related to the origin of the new species as part of a group of troglobitic forms originated from marine interstitial ancestors that colonized the freshwater caves. A key for the identification of all the known species of Stygonitocrellais also provided.     

Finite element analysis of the cranium: Validity,sensitivity and future directions

Finite element analysis (FEA) is increasingly applied in skeletal biomechanical research in general, and in fossil studies in particular. Underlying such studies is the principle that FEA provides results that approximate reality. This paper provides further understanding of the reliability of FEA by presenting a validation study in which the deformations experienced by a real cadaveric human cranium are compared to those of an FE model of that cranium under equivalent simulated loading. Furthermore, model sensitivity to simplifications in segmentation and material properties is also assessed. Our results show that absolute deformations are not accurately predicted, but the distribution of the regions of relatively high and low strains, and so the modes of global deformation, are reasonably approximated.     

Phylogeographic studies of plants in China: Advances in the past and directions in the future

<正>Phylogeography has been one major focus of evolutionary biology in recent years,with many important advances in Chinese species.In this issue,we collected 11 phylogeographic studies of plants by Chinese laboratories. We further synthesized the main findings and patterns emerging from these and previous phylogeographic studies in China and asked where phylogeographic research should be directed in the coming years.Numerous examples have shown that phylogeographic patterns in China did not show an expected expansion-contraction pattern at large scale, mirroring the geological records showing that no unified ice sheet had developed in China during the Quaternary Period.Instead,regional expansions and intraspecific divergences are very common in most studied species during the Quaternary oscillations.Different intraspecific lineages or alleles(haplotypes) were detected in multiple localized refugia,from where regional or local expansions are likely to have started.Hybridizations and introgressions are frequent between intraspecific lineages or between different species.We also reviewed computational methods for phylogeographic analyses.Despite the great progress made in recent years,there remains much to discover about the spatial-temporal dimensions and underlying speciation mechanisms of Chinese plants.Phylogeographic studies represent a key knot that connects the genus phylogeny(macroevolution) and speciation and adaptation (microevolution).Therefore,we advocate that:(i) phylogeographic studies of plants in China should be directed to the closely related species or a monophyletic group(for example,a genus or a section) in the coming years;and (ii) population genetic data based on direct sequencing multiple loci,especially those from nuclear genome and statistical tests should be widely adopted and enforced.The recovered intraspecific divergences and phylogeographic patterns of multiple-species may allow us to better understand the high plant diversity in China and set up concrete hypotheses for studying plant speciation and diversification mechanisms in this region.     

Sperm ultrastructure, morphometry, and abnormal morphology in American black bears (Ursus americanus)

The objective of this study was to describe sperm ultrastructure, morphometry, and abnormal morphology in American black bears. Electroejaculation was successful in 53.8% (7/13) of the attempts, but urine contamination was common. Epididymal sperm samples were also obtained from five bears. Sperm had a paddle-like head shape and the ultrastructure was similar to that of most other mammals. The most striking particularity of black bear sperm ultrastructure was a tightening of the nucleus in the equatorial region. Although the differences were not significant in all bears, the overall decrease in sperm nucleus dimensions during transport from the caput epididymis to the cauda suggested increasing compaction of the nucleus during maturation. For ejaculated sperm, nucleus length, width, and base width were 4.9, 3.7, and 1.8 μm, respectively, whereas sperm head length, width, and base width were 6.6, 4.8, and 2.3 μm, and midpiece, tail (including midpiece), and total sperm lengths were 9.8, 68.8, and 75.3 μm. Evaluation of sperm cytoplasmic droplets in the epididymis revealed that proximal droplets start migrating toward a distal position in the caput epididymis and that the process was mostly completed by the time sperm reached the cauda epididymis. The proportion of morphologically normal sperm in the ejaculate was 35.6%; the most prevalent sperm defects were distal cytoplasmic droplets and bent/coiled tails. The morphology of abnormal sperm and the underlying ultrastructural defects were similar to that in other large domestic animals thus suggesting similar underlying pathogenesis of specific sperm defects and similar effects on fertility.     

Variation in morphology and morphometrics of eggs of Culex quinquefasciatus mosquitoes from different ecological regions of India

Variation in egg surface morphology and morphometrics of Culex quinquefasciatus mosquitoes of the Jodhpur, Bikaner, Jamnagar, and Bathinda strains were correlated with geographical distribution in different ecological regions of India. We report the geographic variation in Cx. quinquefasciatus based on 44 attributes of micropylar and conical‐shaped regions of eggs, including micropylar apparatus (corolla, disc, and mound), micropylar tubercles, and the exochorionic tubercle, pores, and network in anterior, middle, and posterior regions. No remarkable differences were observed in the surface morphology of eggs of these strains except the absence of small tubercles in the anterior and middle region of the JMN strain. However, a statistical analysis indicated significant morphometric variations in 66% of the attributes of the eggs. The cluster analysis of all egg attributes showed that the JD, BKN, and BTH strains are closer to each other than the JMN strain. The positive correlation (r = 0.95) also indicated an effect of geographical distribution on morphometry of various egg attributes of these strains. The present study suggests that ecological variation may have affected the morphometric attributes of the egg of four strains of Cx. quinquefasciatus from different geographical areas.     

TNF family members and malaria: Old observations, new insights and future directions

Tumor necrosis factor (TNF) has long been recognized to promote malaria parasite killing, but also to contribute to the development of severe malaria disease. The precise molecular mechanisms that influence these different outcomes in malaria patients are not well understood, but the virulence and drug-resistance phenotype of malaria parasites and the genetic background and age of patients are likely to be important determinants. In the past few years, important roles for other TNF family members in host immune responses to malaria parasites and the induction of disease pathology have been discovered. In this review, we will summarize these more recent findings and highlight major gaps in our current knowledge. We will also discuss future research strategies that may allow us to better understand the sometimes subtle and intricate effects of TNF family molecules during malaria infection.     

Postcranial variations in late Epigravettian and Neolithic human remains from Arene Candide cave (Liguria,Italy)

Late Epigravettian postcranial human remains from the Arene Candide cave (Finale Ligure, Savona, Italy) were compared with the Neolithic sample found in the upper levels of the same site. Data on length, diaphyseal circumference and diameter of clavicle, humerus, radius, femur and tibia were collected from male specimens having all these bones. The Epigravettian sample is characterized by significantly greater tibial length, robustness and platycnemia, significantly lower circumferences in the upper limb bones and the clavicle, and a high degree of asymmetry. Variations observed in lower limb bones are those expected on the basis of the different functional requirements of a hunting and gathering economy compared to a more sedentary, food producing economy. Differences in the upper limb bones and the clavicle are less explicable. However, considering that in spite of a more slender structure, the Epigravettian bones show evidence of vigorous use, variation in upper limb could result from qualitatively different involvements.     

Deladenus valveus n. sp. (Nematoda: Neotylenchidae) in Dunnage Wood from South Korea Intercepted in Ningbo,China

Deladenus valveus n. sp., isolated from packaging wood originated from South Korea and intercepted in Ningbo, P.R. China, is described and illustrated. Both mycetophagous and infective forms were recovered and are described. The new species D. valveus n. sp. resembles other Deladenus species in which the excretory pore is situated anterior to the hemizonid: in mycetophagous females, the excretory pore is 59 to 74 μm from the anterior end and 37 to 54 μm anterior to the hemizonid. The new species is characterized by the presence of a distinct valve at the esophago-intestinal junction in mycetophagous females and by a degenerate esophagus in mycetophagous males, both of which characters are reported for the first time in a species of Deladenus.     

Home ranges in adult Scandinavian brown bears (Ursus arctos): effect of mass, sex, reproductive category, population density and habitat type

Annual home-range size indices for 36 male and 52 female adult brown bears Ursus arctos in two study areas in central and northern Scandinavia were estimated to evaluate factors believed to influence home-range size. Male home ranges were larger than home ranges of lone females after controlling for the sexual size dimorphism acting on metabolic needs. Further, home ranges of females with cubs were smaller than home ranges of lone females and females with yearlings. Thus, differences in metabolic need were not able to explain the variation in range size among females of different reproductive categories or between males and females, suggesting roaming behaviour of males in this promiscuous species. Home-range size in both males and females was inversely related to population density along a density gradient that was not linked to food availability. This contradicts the hypothesis that females use the minimum areas that sustain their energy requirements. However, on a large geographical scale a negative relationship between range size and food availability was evident. The annual home ranges in inland boreal environments in Scandinavia are the largest reported for brown bears in Eurasia, and similar to those in inland boreal and montane environments in North America.