Body size of the red fox Vulpes vulpes in Spain: the effect of agriculture

It is well recognized that differences in environmental selection pressures among populations can generate phenotypic divergence in a suite of morphological characteristics and associated life history traits. Previous analysis of mitochondrial DNA and body size have suggested that Black Bears (Ursus americanus) inhabiting the island of Newfoundland represent a different subspecies or ecotype from mainland populations. Assuming that body size covaries positively with skull size, we predicted that skull size would be greater for bears on the island than the mainland, and the distribution of size-related shape components in multivariate space should show a distinct separation between Newfoundland and mainland populations. Measurements of 1080 specimens from Newfoundland, Alberta, New York, and Quebec did not provide unequivocal support for our prediction that skull size in Newfoundland bears would be larger than bears from the mainland populations. After removing ontogenetic effects of skull size, between-population variation in skull shape was greater in females than males, and the analysis significantly separated Newfoundland bears from mainland populations. Explanations for this pattern are numerous, but currently remain hypothetical. Limited covariation between skull size and body size suggests that genetic traits regulating the size of Black Bear skulls are more heritable (i.e., less influenced by environmental selection pressures) than characteristics affecting body size. We hypothesize that if gape size does not limit prey size in solitary terrestrial carnivores, large degrees of among-population variation in body size should be coupled with little covariation in skull size. In general, sexual dimorphism in skull size and shape was marginal for the phenotypic characters measured in our study. We believe that sexual dimorphism in skull size in Black Bears is primarily driven by intrasexual selection in males for increased gape size display, while similarity in skull shape between sexes is associated with the constraints of a temporally-selective, but similar diet.

Observations on variation in skull size of three mammals in Israel during the 20th century

We studied geographical and temporal body size trends among 169 adult museum specimens of the Eurasian otter (Lutra lutra) collected in Sweden between 1962 and 2008, whose sex, year of collection, and locality were known. Skull size and body mass increased significantly in relation to the year of collection, and skull size (but not body mass) was significantly and negatively related to latitude, contrasting Bergmann's rule and the trend found for Norwegian otters. Latitudinal differences in body size between the two countries may be due to differences in food availability. The temporal increase in body size among Swedish otters resembled that observed for Norway otters, though Swedish otters are smaller with respect to their Norwegian counterparts. Latitude and year represent a combination of environmental factors, including ambient temperature in the year of collection as well as the number of days of ice coverage. We replaced the above factors with mean annual temperature or the number of days of ice coverage, and found that each of these factors explains a similar proportion of the variation in body size as did latitude and year. We hypothesize that this temporal increase in body size is related to a combination of factors, including reduced energy expenditure resulting from increasing ambient temperature, and increased food availability from longer ice-free periods.

Using museum specimens, we studied temporal changes in skull size in two species of Japanese rodents, the large Japanese field mouse (Apodemus speciosus) and Pratt's vole (Eothenomys smithii= E. kageus) during the 20th century. We used the greatest length of the skull (GTL), zygomatic breadth (ZB), narrowest width of the skull across the interorbital region (IC) and the length of the upper cheek teeth row (M) as indicators to such changes. We found that GTL and ZB (but not IC and M) increased during the study period in mice, and that IC and M (but not GTL and ZB) increased marginally in voles. We attribute these changes to elevated ambient minimal temperatures, which increased food availability and energy savings for the mice, and required diet change in the voles.

Understanding the spatial distribution of phenotypes and their association with local environmental conditions can provide important insights into the evolutionary history and ecological dynamics of species. Geographical variation in the skull size of the Artibeus lituratus complex was explored to evaluate the association between morphological traits and habitat-specific environmental conditions in the mainland populations of Middle and South America. We performed a principal component analysis based on 390 museum specimens using 17 cranial and mandibular measurements to explore the overall morphometric variation in our sample. Additionally, we used the information from 19 bioclimatic variables from 127 collecting localities to assess the extent of variation in environmental space across our study area. A canonical correlation analysis performed to evaluate the association between morphological and environmental variables indicated a high correlation between morphology and environment. Seasonality was correlated with skull size (canonical r = 0.7). Specifically, skull size in the A. lituratus complex increases as the amount of precipitation during the driest season increases and as inter-annual precipitation variability decreases. This result supports the hypothesis that environmental pressures are at least partially responsible for the skull size differences observed. Moreover, the nonrandom distribution of individuals with different skull sizes suggests that environmental filtering plays a role in determining the geographical distribution of morphological variants.

Using museum material, we studied temporal changes in skull size of 185 red foxes (Vulpes vulpes Linnaeus, 1758) and 71 Eurasian badgers (Meles meles Linnaeus, 1758) collected in Austria between 1866 and 2007. Four measurements were taken and combined into principal components by means of a principal components analysis. Akaike's information criterion models indicated that skull size of the red fox and the Eurasian badger is positively related to the year of collection and negatively to altitude but not to latitude or longitude. However, for the Eurasian badger, the relationship between skull size and year is only on the verge of significance. We suggest that the increase in skull size with year is a result of improved food availability during the 20th Century from man-made resources such as agricultural produce and garbage, and the decrease in skull size with altitude is possibly the result of a parallel decrease in primary productivity. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●●, ●●–●●.

Using a geometric morphometric approach, we explored the variation in skull size and skull shape in banded newts (genus Ommatotriton). The genus Ommatotriton is represented by two allopatric, genetically well‐defined species: Ommatotriton ophryticus and O. vittatus. Within each species, two subspecies have been recognised. The samples used in this study cover the geographical and genetic variation within each species. We found statistically significant variation in skull size between species and among populations within species. When corrected for size, there was no significant variation in shape between species. Our results indicate that the variation in skull shape within the genus Ommatotriton is almost entirely due to size‐dependent, allometric shape changes. The exception is the shape of the ventral skull in males. Males of O. ophryticus and O. vittatus significantly diverge in the shape of the ventral cranium. The ventral skull, more precisely the upper jaw and palate, is directly functionally related to feeding. In general, our results indicate that allometry is a significant factor in the morphological variation of banded newts. However, the divergence in the ventral skull shape of males indicates that sexual selection and niche partitioning may have influenced the evolution of skull shape in these newts.

The lynx Lynx canadensis is a common predator in the boreal forests of North America. Its population fluctuates during a 9- to 11-year cycle in synchrony with the population size of its main prey, the snowshoe hare Lepus americanus. Using adult museum specimens, we studied changes in skull (and hence body) size of the lynx in Alaska during the second half of the 20th century. The population cycle in Alaska averaged 9 years, similar to that reported in the neighbouring Yukon. Using harvest data of lynx as an estimate of population size, we found that skull size was negatively related to population size. This relationship was strongest not for the population density in the year of death (X), but for year X-3, a carry-over effect from the first year (or years) of life, indicating that conditions during the fast-growth years are determining body size. We suggest that the density-dependent effect is probably due to changes in food supply, either resulting from the adverse effects of competition or a possible diminished availability of food. Two skull parameters decreased significantly during the second half of the 20th century. We do not know the cause for the year effect and suggest that it might be due to a long-term change in the availability of prey. Canine size did not change during the study period, probably an indication that snowshoe hares maintained their status as the main prey of the lynx throughout the study period.

Amphisabenia is a group of squamates adapted for a fossorial lifestyle. The skull is the animal's the main digging tool, and can present one of four principal shapes. The shovel-headed shape is considered to be the most specialized for digging. The South American genus Leposternon presents a shovel-headed morphotype, and is widely distributed on this continent. The general shovel-headed skull pattern may vary considerably, even within the same genus, and we hypothesized that this variation may be influenced primarily by body size and geographical factors. This study investigated the variation in skull size and shape among five Leposternon species, and examined the potential relationship between this variation and the size of the specimens and bioclimatic variables, through a geometric morphometric approach. Significant morphological variation was found among the species, and was also related systematically to body size and the geographical distribution of the specimens. As even subtle differences in the skull size or shape may represent significant modification in bite force and digging capacity and digging speed, the cranial variation found among the Leposternon species and specimens may have a direct influence on their diet and locomotor performance. Our results, together with direct observations of some of these species, suggest that shovel-headed amphisbaenians may be able to penetrate different soil types under a range of climatic conditions, especially considering the ample, but often sympatric distribution of the species studied here.

Changes in body size inversely related to ambient temperatures have been described in woodrats (Neotoma) over time scales ranging from decades to millennia. However, climate-mediated variation in other traits has not been evaluated, and the effects of precipitation have been overlooked. We assessed variation in skull morphology among bushy-tailed woodrats (Neotoma cinerea) over two sampling transects spanning coastal rainforest and interior desert environments to determine whether skull morphology varied with climate. We also tested whether previously described size-temperature relationships could be generalized to our study populations. In both transects, linear measurements of functionally significant traits differed between coastal and interior populations. Geometric morphometric analyses of shape confirmed some of those differences and revealed additional patterns of skull variation. Variation in some linear measurements, including body size, was predicted by climate. However, body and skull size, as well as measurements of skull components, displayed varying responses. Although longitudinal patterns of body size variation supported Bergmann’s rule, skull size variation was only weakly associated with climate. The strongest phenotypic responses to climate were those of auditory, dental, and palatal skull traits. Altogether, our findings suggest that geographic variation in temperature and precipitation mediated selective heterogeneity and plasticity in skull traits associated with food processing and sensory organs in N. cinerea. This was consistent with our expectation of resource-dependent phenotypic variation among populations in environments with highly contrasting climatic regimes.

Commercial whaling peaked between the 1700s and 1800s, leading to declines in whale populations worldwide. While it is well constrained that baleen whales have shrunk in body size over the past centuries - likely due to whaling pressure and climate change – smaller toothed whale species, which were not the primary targets during the whaling era, have been largely understudied in this regard.However, one of the major challenges in the face of global change and increasing anthropogenic influence is predicting population declines in order to establish suitable conservation strategies before a collapse can take place. Tracking declines in body sizes over large temporal scales has proven to be a reliable indicator preceding such population declines and collapses.In this study we investigate body sizes of the harbour porpoise (Phocoena phocoena), which is one of the smallest toothed cetacean species and a common inhabitant of European waters. We use skull sizes as proxies for estimating body sizes of individuals collected from Scotland, England and the Netherlands, with collection dates spanning from the 17th century to the present. The aim of this study is to assess whether a decline in harbour porpoise body sizes over time and additionally, if possible morphospecies from different regions of the North Sea, can be identified.

The upstream binding factor 1 (UBF1) is one of the proteins in a complex that regulates the activity of RNA polymerase I, which controls the rate of ribosomal RNA (rRNA) synthesis. We have shown previously that insulin receptor substrate-1 (IRS-1) can translocate to the nuclei and nucleoli of cells and bind UBF1. We report here that activation of the type I insulin-like growth factor receptor (IGF-IR) by IGF-I increases transcription from the ribosomal DNA (rDNA) promoter in both myeloid cells and mouse fibroblasts. The increased activity of the rDNA promoter is accompanied by increased phosphorylation of UBF1, a requirement for UBF1 activation. Phosphorylation occurs on a number of UBF1 peptides, most prominently on the highly acidic, serine-rich C terminus. In myeloid cells (but not in mouse embryo fibroblasts) IRS-1 signaling stabilizes the levels of UBF1 protein. These findings demonstrate that IGF-IR signaling can increase the activity of UBF1 and transcription from the rDNA promoter, providing one explanation for the reported effects of the IGF/IRS-1 axis on cell and body size in animals and cells in culture.

Little is known about the large‐scale evolutionary patterns of skull size relative to body size, and the possible drivers behind these patterns, in Archosauromorpha. For example, the large skulls of erythrosuchids, a group of non‐archosaurian archosauromorphs from the Early and Middle Triassic, and of theropod dinosaurs are regarded as convergent adaptations for hypercarnivory. However, few investigations have explicitly tested whether erythrosuchid and theropod skulls are indeed disproportionately large for their body size, and whether this trend is driven by hypercarnivory. Here, we investigate archosauromorph relative skull size evolution, examining the scaling relationships between skull and body size of Palaeozoic and Mesozoic archosauromorphs using a robust phylogenetic framework and assessing the influence of potential drivers, such as taxonomy, diet, locomotory mode and inhabited biotope. Our results show that archosauromorph relative skull sizes are largely determined by phylogeny and that the other drivers have much weaker levels of influence. We find negative allometric scaling of skull size with respect to body size when all studied archosauromorphs are analysed. Within specific groups, skull size scales with positive allometry in non‐archosaurian archosauromorphs and, interestingly, scales isometrically in theropods. Ancestral reconstructions of skull–femur size ratio reveal a disproportionately large skull at the base of Erythrosuchidae and proportionately sized skulls at the bases of Theropoda, Carnosauria and Tyrannosauroidea. Relative skull sizes of erythrosuchids and theropods are therefore distinct from each other, indicating that disproportionately large skulls are not a prerequisite for hypercarnivory in archosauromorphs, and that erythrosuchids exhibit a bauplan unique among terrestrial Mesozoic carnivores.

Isoodon obesulus exhibits geographic variation in body size and shape, which appears to be adaptive. The geographic range of this species is declining, so the presence of adaptive divergence is of concern for the conservation of this species, both in the long term (loss of diversity decreasing evolutionary potential) and short term (choice of source populations for translocations). In this study, skulls of I. obesulus , both recent (animals alive within the last 100 years) and fossil (a few thousand years old), were examined and a range of measurements obtained. Comparisons were made between the two data sets to see whether skull morphology has changed over this relatively short period. Such a change may indicate rapid evolution of these characters and therefore the potential for fast regeneration of any lost geographic variation. Fossil skulls were smaller than their recent counterparts, had shorter ‘snouts’ relative to skull width and depth, and displayed no geographic variation in size and shape, whereas recent skulls were geographically differentiated. Because of the apparent rapid evolution in these characters, the implications of adaptive variation in size and shape in I. obesulus with regard to its conservation may be strictly short term, since any geographic variation lost may be quickly recovered if suitable conditions exist.

New World monkeys (NWM) display substantial variation (two orders of magnitude) in body size. Despite this, variation in skull size and associated shape show a conserved allometric relationship, both within and between genera. Maximum likelihood estimates of quantitative ancestral states were used to compare the direction of morphological differentiation with the phenotypic (pmax) and genetic (gmax) lines of least evolutionary resistance (LLER). Diversification in NWM skulls occurred principally along the LLER defined by size variation. We also obtained measures of morphological amount and pace of change using our skull data together with published genetic distances to test whether the LLER influenced the amount and pace of diversification. Moreover, data on an ecological factor (diet) was obtained from the literature and used to test the association of this niche-related measure with the morphological diversification. Two strategies were used to test the association of LLER with the morphological and dietary amount and pace of change, one focusing on both contemporary genera and maximum likelihood reconstructed ancestors and the other using only the 16 contemporary genera in a phylogenetic comparative analysis. Our results suggest that the LLER influenced the path, amount, and pace of morphological change. Evolution also occurred away from the LLER in some taxa but this occurred at a slower pace and resulted in a relatively low amount of morphological change. We found that longer branch lengths (time) are associated with larger differences in pmax orientation. However, on a macroevolutionary scale there is no such trend. Diet is consistently associated with both absolute size differences and morphological integration patterns, and we suggest that this ecological factor might be driving adaptive radiation in NWM. Invasion of diet-based adaptive zones involves changes in absolute size, due to metabolic and foraging constraints, resulting in simple allometric skull diversification along the LLER. While it is clear that evolutionary change occurred along the LLER, it is not clear whether this macroevolutionary pattern results from a conservation of within-population genetic covariance patterns or long-term adaptation along a size dimension or whether both constraints and selection were inextricably involved.