The Neanderthal "chignon": variation, integration, and homology

The occipital bun ("chignon") is cited widely as a Neanderthal derived trait. It encompasses the posterior projection/convexity of the occipital squama and is associated with lambdoid flattening on the parietal. A 'hemibun' in some Upper Paleolithic Europeans is thought by some authors to indicate interbreeding between Neanderthals and early modern Europeans. However, 'bunning' is difficult to measure, and the term has been applied to a range of morphological patterns. Furthermore, its usefulness in phylogenetic reconstruction and its homologous status across modern and fossil humans have been disputed. We present a geometric morphometric study that quantitatively evaluates the chignon, assesses its usefulness in separating Neanderthals from modern humans, and its degree of similarity to Upper Paleolithic 'hemibuns.' We measured the three-dimensional coordinates of closely spaced points along the midsagittal plane from bregma to inion and of anatomical landmarks in a large series of recent human crania and several Middle and Late Pleistocene European and African fossils. These coordinate data were processed using the techniques of geometric morphometrics and analyzed with relative warps, canonical variates, and singular warps. Our results show no separation between Neanderthals and modern humans, including early modern Europeans, when the shape of the occipital plane midsagittal-profile is considered alone. On the other hand, Neanderthals are well separated from both recent and fossil modern humans when information about the occipital's relative position and relative size are also included. Furthermore, the occurrence of a highly convex and posteriorly projecting midline occipital profile (interpreted as the occipital bun) is highly correlated (>0.8) with a flat parietal midsagittal profile and with antero-superiorly positioned temporal bones across both our recent and our fossil human samples. We conclude that the shape of the occipital profile alone should not be considered an independent trait, as it is very tightly integrated with braincase shape. Our analysis does not support differences in integration of the posterior midsagittal profile and the cranial base in Pleistocene and recent humans.     

Cranial integration in Homo: singular warps analysis of the midsagittal plane in ontogeny and evolution

This study addresses some enduring issues of ontogenetic and evolutionary integration in the form of the hominid cranium. Our sample consists of 38 crania: 20 modern adult Homo sapiens, 14 sub-adult H. sapiens, and four archaic Homo. All specimens were CT-scanned except for two infant H. sapiens, who were imaged by MR instead. For each specimen 84 landmarks and semi-landmarks were located on the midsagittal plane and converted to Procrustes shape coordinates. Integration was quantified by the method of singular warps, a new geometric-statistical approach to visualizing correlations among regions. The two classic patterns of integration, evolutionary and ontogenetic, were jointly explored by comparing analyses of overlapping subsamples that span ranges of different hypothetical factors. Evolutionary integration is expressed in the subsample of 24 adult Homo, and ontogenetic integration in the subsample of 34 H. sapiens. In this data set, vault, cranial base, and face show striking and localized patterns of covariation over ontogeny, similar but not identical to the patterns seen over evolution. The principal differences between ontogeny and phylogeny pertain to the cranial base. There is also a component of cranial length to height ratio not reducible to either process. Our methodology allows a separation of these independent processes (and their impact on cranial shape) that conventional methods have not found.     

Form changes in the sea bass, Dicentrarchus labrax (Moronidae: Teleostei), after acclimation to freshwater: an analysis using shape coordinates

Synopsis Morphological changes in the sea bass, Dicentrarchus labrax (Perciformes: Moronidae), were investigated after an experimental acclimation trial to freshwater. The sea bass is an euryhaline species occurring in the Mediterranean and west Atlantic from 30° N to 55° N. Part of the offspring of a pool of breeders was acclimated to freshwater at 9 months of age while maintaining the original stock in marine water. The effect of acclimation to freshwater over the entire form of the fish was studied through geometric morphometrics (shape coordinates). Changes in the form were shown graphically as landmark displacements though age classes 7,12, 15, 19 and 24 months and were discussed in the light of integrated growth. A significantly faster growth in freshwater was detected. Shape coordinates analyzed through multivariate statistics show that significant differences in shape arise after acclimation to freshwater. Moreover, the uniform component of shape change reveals a major effect of stretching or compression perpendicular to the body axis which is common and conservative in both trials. Differences are discussed in terms of selection and ecophenotypism.     

Discrimination of the sibling species Apodemus flavicollis and A. sylvaticus (Rodentia,Muridae)

Karyotyping and several molecular methods have allowed successful identification of two morphologically similar wide-ranging Western Palearctic species, the yellow-necked field mouse Apodemus flavicollis (Melchior, 1934) and the long-tailed wood mouse A. sylvaticus (Linnaeus, 1758), but reliable species diagnosis on the basis of morphometric characters is particularly problematic. Although they are easily morphologically distinguishable in Central and Northern Europe, this is not the case in southern parts of their distribution areas. Despite that, we have successfully discriminated A. flavicollis and A. sylvaticus from Serbia (Southern Europe) using geometric and traditional morphometric methods on a data set for ventral crania of specimens previously genotyped by the Inter Simple Sequence Repeat-PCR (ISSR-PCR). Discrimination power of applied approaches was more or less similar. The majority of our results were consistent with those obtained for specimens collected across the Czech Republic (Central Europe). Morphological differences observed herein, as well as those already reported between A. flavicollis and A. sylvaticus from the central and northern parts of their distribution areas, could be the outcome of their biology, i.e. ecological discrepancies, different assumed evolutionary scenarios considering biogeography, phylogeny, history and ontogeny.     

Left–right asymmetries and shape analysis on Ceroglossus chilensis (Coleoptera: Carabidae)

Bilateral symmetry is widespread in animal kingdom, however most animal can deviate from expected symmetry and manifest some kind of asymmetries. Fluctuating asymmetry is considered as a tool for valuating developmental instability, whereas directional asymmetry is inherited and could be used for evaluating evolutionary development. We use the method of geometric morphometrics to analyze left/right asymmetries in the whole body, in two sites and totally six populations of Ceroglossus chilensis with the aim to infer and explain morphological disparities between populations and sexes in this species. In all individuals analyzed we found both fluctuating asymmetry and directional asymmetry for size and shape variation components, and a high sexual dimorphism. Moreover a high morphological variability between the two sites emerged as well. Differences in diet could influence the expression of morphological variation and simultaneously affect body sides, and therefore contribute to the symmetric component of variation. Moreover differences emerged between two sites could be a consequence of isolation and fragmentation, rather than a response to local environmental differences between sampling sites.     

On the comparison of the strength of morphological integration across morphometric datasets

Evolutionary morphologists frequently wish to understand the extent to which organisms are integrated, and whether the strength of morphological integration among subsets of phenotypic variables differ among taxa or other groups. However, comparisons of the strength of integration across datasets are difficult, in part because the summary measures that characterize these patterns (RV coefficient and r_PLS) are dependent both on sample size and on the number of variables. As a solution to this issue, we propose a standardized test statistic (a z‐score) for measuring the degree of morphological integration between sets of variables. The approach is based on a partial least squares analysis of trait covariation, and its permutation‐based sampling distribution. Under the null hypothesis of a random association of variables, the method displays a constant expected value and confidence intervals for datasets of differing sample sizes and variable number, thereby providing a consistent measure of integration suitable for comparisons across datasets. A two‐sample test is also proposed to statistically determine whether levels of integration differ between datasets, and an empirical example examining cranial shape integration in Mediterranean wall lizards illustrates its use. Some extensions of the procedure are also discussed.     

Shape and size variations in the cranium of elephant-shrews: a morphometric contribution to a phylogenetic debate

A geometric morphometric analysis was carried out on the crania of 13 species of elephant-shrews (Macroscelidea), a group of African mammals whose phylogeny is still debated. The material examined consisted of 313 crania and included all the genera of Macroscelididae, the unique family recognized by taxonomists. The results obtained from the analysis of the cranium shape and size, either from dorsal or lateral view, were very similar. The first one appeared more reliable because of the higher number of intersection points fixed between the cranial sutures. All the cranial features that distinguished the genus Rhynchocyon were a consequence of the extreme enlargement of frontal bones. Instead, within the subfamily Macroscelidinae, the differences between genera were based on modification involving other bones, mainly mastoids and nasals, as shown by the deformation grids. A cluster analysis confirmed the traditional subdivision in two subfamilies (Rhynchocyoninae and Macroscelidinae) but suggested a different relationship among the recognized genera belonging to Macroscelidinae. Our results are congruent with data obtained from previous biochemical research and support the traditional subdivision in two subfamilies, the monophily of the genus Elephantulus and its closeness with Petrodromus, relating to their similar cranium shapes. The latter presumably is a case of gigantism as adaptation to forest habitats. Further studies on all the species of Elephantulus could provide new evidence for assessing the relationships within this clade, including Macroscelides that by the present analysis appeared as a well-distinguished taxonomic entity.     

The Vindija Neanderthal scapular glenoid fossa: comparative shape analysis suggests evo-devo changes among Neanderthals

Although the shape of the scapular glenoid fossa (SGF) may be influenced by epigenetic and developmental factors, there appears to be strong genetic control over its overall form, such that variation within and between hominin taxa in SGF shape may contain information about their evolutionary histories. Here we present the results of a geometric morphometric study of the SGF of the Neanderthal Vi-209 from Vindjia Cave (Croatia), relative to samples of Plio-Pleistocene, later Pleistocene, and recent hominins. Variation in overall SGF shape follows a chronological trend from the plesiomorphic condition seen in Australopithecus to modern humans, with pre-modern species of the genus Homo exhibiting intermediate morphologies. Change in body size across this temporal series is not linearly directional, which argues against static allometry as an explanation. However, life history and developmental rates change directionally across the series, suggesting an ontogenetic effect on the observed changes in shape (ontogenetic allometry). Within this framework, the morphospace occupied by the Neanderthals exhibits a discontinuous distribution. The Vindija SGF and those of the later Near Eastern Neanderthals (Kebara and Shanidar) approach the modern condition and are somewhat segregated from both northwestern European (Neandertal and La Ferrassie) and early Mediterranean Neanderthals (Krapina and Tabun). Although more than one scenario may account for the pattern seen in the Neanderthals, the data is consistent with palaeogenetic evidence suggesting low levels of gene flow between Neanderthals and modern humans in the Near East after ca. 120-100 ka (thousands of years ago) (with subsequent introgression of modern human alleles into eastern and central Europe). Thus, in keeping with previous analyses that document some modern human features in the Vindija Neanderthals, the Vindija G₃ sample should not be seen as representative of ‘classic’ - that is, unadmixed, pre-contact - Neanderthal morphology.     

白尾鼹(鼹科: 哺乳纲)下颌骨几何形态测量分析及地理分化研究

白尾鼹属为东洋界单型属,主要分布于中国西南山地、印度阿萨姆地区、孟加拉国、缅甸北部和老挝等地。由于白尾鼹分布区范围内地形、气候和生境十分复杂,很可能导致物种的地理分化。为此我们查看了分布区之内18 个地区的49 号标本的下颌骨,并基于几何形态测量的方法进行分析比较。结果显示所有的标本按照地理分布可分为3 组(滇东北、横断山南段以及印度)。3 组标本间的形态差异主要表现在水平枝上,其中下臼齿齿列的长度、下颌骨水平枝下缘的弯曲度以及水平枝中部高度差异最为明显,而上升枝对3 个地理居群的区分能力较弱。其中滇东北标本的下颌骨最为平直,下臼齿列长度最短;而横断山居群的下颌骨弯曲度最大,下臼齿列最长。根据以上结果,我们推测3 个下颌骨形态有明显差异的地理居群可能代表了3 个不同的地理亚种或是不同的种。但这仍需要结合上颌骨的形态差异和遗传结构分析加以证实。     

Morphological cranial diversity contributes to phylogeny in soft-furred sengis (Afrotheria, Macroscelidea)

Despite the well-supported Macroscelidea phylogeny proposed at the end of the 1960s, several systematic arrangements have been suggested in the last 20 years, raising doubts about the phylogeny of the Macroscelidinae; sengi inter-specific relationships are still debated to this day. The main issue of concern involves the supposed Elephantulus diphyly. To solve this persisting debate about sengi phylogeny, we examined the cranium ventral surface of 13 species using geometric morphometric techniques and neighbour-joining algorithms. This study supported the idea that the ventral side of the sengi cranium has the potential to provide important signals for reconstructing the Macroscelidea phylogeny. The phylogenetic signals seemed to differentiate between two major clades in the sengi radiation. In the first clade, the two monospecific genera (Petrodromus and Macroscelides), the two African Horn species (Elephantulus revoilii and E. rufescens), and the only North African species (E. rozeti) were clustered together. The second clade includes the remnant south-central African Elephantulus species. Our results were in agreement with both mitochondrial and nuclear data, confirmed that there is no Elephantulus monophyly and highlighted the close relationship between Petrodromus and E. rozeti. It appears that all the soft-furred sengi species are organised in two evolutionary lines: an old monophyletic clade, comprising only Elephantulus species, and a new polyphyletic clade, including P. tetradactylus, M. proboscideus, and E. rozeti. This requires a taxonomic and nomenclatural rearrangement within Macroscelidinae, where the phylogenetic position of the remnant 4 (of 12) Elephantulus species has yet to be fully defined.     

Evolution of extreme ontogenetic allometric diversity and heterochrony in pythons,a clade of giant and dwarf snakes

Ontogenetic allometry, how species change with size through their lives, and heterochony, a decoupling between shape, size, and age, are major contributors to biological diversity. However, macroevolutionary allometric and heterochronic trends remain poorly understood because previous studies have focused on small groups of closely related species. Here, we focus on testing hypotheses about the evolution of allometry and how allometry and heterochrony drive morphological diversification at the level of an entire species‐rich and diverse clade. Pythons are a useful system due to their remarkably diverse and well‐adapted phenotypes and extreme size disparity. We collected detailed phenotype data on 40 of the 44 species of python from 1191 specimens. We used a suite of analyses to test for shifts in allometric trajectories that modify morphological diversity. Heterochrony is the main driver of initial divergence within python clades, and shifts in the slopes of allometric trajectories make exploration of novel phenotypes possible later in divergence history. We found that allometric coefficients are highly evolvable and there is an association between ontogenetic allometry and ecology, suggesting that allometry is both labile and adaptive rather than a constraint on possible phenotypes.     

Cranial and mandibular morphology of Middle Pleistocene cave bears (Ursus deningeri): implications for diet and evolution

ABSTRACT

Deninger’s bears (Ursus deningeri) have been studied less frequently than Ursus spelaeus s.l. Our objective is to present, for the first time, an analysis of the skull shape of U. deningeri.

Bear crania and mandibles were digitised with a Microscribe or CT-scanned and the surface models subsequently landmarked. The landmarks were chosen based on a compromise between functional morphology and sample size.

Results show that U. deningeri and U. spelaeus mandibles display very similar morphologies and allometric trajectories, both to each other and to Ailuropoda melanoleuca. It is inferred that masticatory adaptations to a herbivorous diet were already present in the Middle Pleistocene. U. deningeri displays a cranial morphology that is similar to that of U. spelaeus when comparing all species, but U. deningeri has a relatively narrower and dorsoventrally lower zygomatic arch than U. spelaeus, although the masticatory signal is less strong in the skull.

We observe intraspecific differences between different populations of U. deningeri, which could parallel the genetic diversity found in U. spelaeus. The intraspecific differences found within U. deningeri may be temporal and/or geographical in nature and could be related to the evolution of the Late Pleistocene cave bear, but this hypothesis remains to be tested.     

华南长江和西江流域鲤科宽鳍鱲和马口鱼不同线粒体DNA谱系之间的体形分化

本文研究了两种鲤科鱼类,即分布于华南长江与西江的宽鳍鱲和马口鱼的不同线粒体DNA谱系之间的体形差异。近期基于线粒体DNA的这两个物种的系统发育分析认为这两个类元内包含多个谱系(宽鳍鱲A-D和马口鱼1-5)。本文采用几何形态度量学的方法研究了这些不同线粒体DNA谱系间是否存在体形方面的差异。在宽鳍鱲的4个不同的线粒体DNA谱系中,有3个被纳入本研究内容。结果显示:在宽鳍鱲的3个线粒体DNA谱系中,2个具有体形的差异。在马口鱼的5个线粒体DNA谱系中,3个被纳入本文的研究。结果显示所有3个谱系均存在体形差异。因此,线粒体DNA的谱系差异很大程度上表现在谱系间的体形差异方面,表明线粒体DNA谱系可能对应于不同的物种。     

华南长江和西江流域鲤科宽鳍鳞和马口鱼不同线粒体DNA谱系之间的体形分化

本文研究了两种鲤科鱼类,即分布于华南长江与西江的宽鳍雠和马口鱼的不同线粒体DNA谱系之间的体形差异。近期基于线粒体DNA的这两个物种的系统发育分析认为这两个类元内包含多个谱系（宽鳍鱲A—D和马口鱼1—5）。本文采用几何形态度量学的方法研究了这些不同线粒体DNA谱系间是否存在体形方面的差异。在宽鳍馘的4个不同的线粒体DNA谱系中,有3个被纳入本研究内容。结果显示：在宽鳍雠的3个线粒体DNA谱系中,2个具有体形的差异。在马口鱼的5个线粒体DNA谱系中,3个被纳入本文的研究。结果显示所有3个谱系均存在体形差异。因此,线粒体DNA的谱系差异很大程度上表现在谱系间的体形差异方面,表明线粒体DNA谱系可能对应于不同的物种。     

Quantitatively comparing morphological trends to environment in the fossil record (Cincinnatian series; Upper Ordovician)

Determining whether morphological trends in fossil species represent evolution within a lineage or lateral shifts in morphologically variable populations through time requires a thorough examination of the details of both morphology and paleoenvironment in time and space. The purpose of this study is to explore at high resolution the relationship between morphology of the trilobite Flexicalymene granulosa and paleoenvironmental conditions in Upper Ordovician deposits of southwestern Ohio and northern Kentucky. This is achieved by using geometric morphometrics to measure high-resolution morphological changes and by using gradient analysis to capture environmental gradients underlying faunal distribution patterns. Quantitatively comparing the outcomes of these two techniques provides an assessment of whether shape changes relates to environment. Results indicate that a significant amount of shape change, seen as an anteromedial movement of the eye region over time, corresponds to ordination scores. This suggests a relationship between certain aspects of morphology and environment. The combination of these quantitative techniques has provided the foundation for determining whether morphological trends within F. granulosa represent evolution or temporal shifts in geographic variation. Future work will involve examining this relationship in greater detail both geographically and stratigraphically.     

A phylogenetic view on skull size and shape variation in the smooth newt (Lissotriton vulgaris,Caudata, Salamandridae)

In this study, we explore skull size and shape variation in the smooth newt, a taxon with substantial morphological differentiation and complex phylogeographic relations. By projecting phylogenies into the morphospace of the skull shape, we explore the variation in and differentiation of this complex morphological structure within a phylogenetic framework. For these analyses, we used a dataset that covers the most southern part of the species’ distribution range, including all conventionally recognized subspecies. The study revealed different patterns of divergence in skull shape between sexes, which is paralleled by intraspecific differentiation. The divergence in dorsal skull shape is concordant with the phylogenetic divergence, as the most diverged clades of the smooth newt (Lissotriton vulgaris kosswigi and Lissotriton vulgaris lantzi) exhibit a skull shape that significantly diverges from the smooth newt’s mean shape configuration. The results of this study also indicate that ventral skull portion, which is more directly related to feeding and foraging, shows higher variation between populations than dorsal skull portion, which appears to be less variable and phylogenetically informative.