Variation in Seasonal Movement and Body Size of Wintering Populations of Black-Headed Gull in Japan

The relationship between environmental gradients and patterns of geographic variation in body size has been a controversial topic for ectothermic organisms globally. To examine whether the patterns that generally hold in more temperate species also hold for tropical ones, we examined the intraspecific body size variation in three species of Neotropical frogs, Dendropsophus minutus, Hypsiboas faber and Physalaemus cuvieri, along different environmental gradients (e.g. temperature, precipitation and topography). We analysed four competing hypotheses: (i) the water availability hypothesis that predicts a negative relationship between body size and precipitation; (ii) the heat balance hypothesis that predicts a negative relationship between body size and temperature; (iii) the topography hypothesis that predicts a negative relationship between body size and altitude; and (iv) the mixed-effect hypothesis that predicts that individuals occurring in wet and cold sites would be larger than individuals occurring in dry and warm sites. The spatial pattern of geographic variation in body size among populations of H. faber was associated with the mixed-effect hypothesis. In localities with low precipitation seasonality and cold conditions, H. faber individuals were larger than in localities with high precipitation seasonality and warm conditions. Variation in the body size of D. minutus was the opposite of that predicted by the heat balance hypothesis. Individuals in localities with high temperatures were larger than in localities with low temperatures. On the other hand, variation in the body size of P. cuvieri was not associated with the variables used in this study. Our results suggest that intraspecific variation in anuran body size is more dependent on species-specific response than on the region (i.e. temperate or tropical) where they occur.

ABSTRACTAim To determine how well variation in median body size of avian assemblages is predicted by (1) the environmental models usually employed in analyses of Bergmann's rule and (2) random sampling from the regional body size frequency distribution. If body size frequency distributions of local assemblages represent a random sample of a regional frequency distribution, then geographical variation in body sizes of assemblages might be a consequence of the determinants of spatial variation in species richness rather than direct influences on body size per se.Location Southern Africa.Methods Median body masses (as a measure of body size) of avian assemblages were calculated for quarter‐degree grid cells across South Africa and Lesotho. The relationship between median body mass and four environmental variables (minimum and maximum monthly temperatures, precipitation and seasonality in the normalized difference vegetation index, as a measure of seasonality in productivity) was examined using general linear models first without taking spatial autocorrelation into account, and then accounting for it by fitting an exponential spatial covariance structure. Model fit was assessed using the Akaike information criterion and Akaike weights. At each species richness value, random assemblages were sampled by either drawing species randomly from the regional body mass frequency distribution, or drawing species from the regional body mass frequency distribution with a probability proportional to their geographical distribution in the area. The ability of randomizations to predict actual body masses was examined using two‐tailed Fisher exact tests.Results Seasonality in productivity was the only environmental variable that remained a significant predictor of body mass variation in spatially explicit models, though the positive relationship was weak. When species richness was included in the models it remained the only significant predictor of size variation. Randomizations predicted median body mass poorly at low species richness, but well at high richness.Main conclusions Environmental models that have previously been proposed explain little of the variation in body mass across avian assemblages in South Africa. However, much of the variation in the median mass of assemblages could be predicted by randomly drawing species from the regional body mass frequency distribution, particularly using randomizations in which all species were drawn from the regional body mass frequency distribution with equal probability and at high species richness values. This outcome emphasizes the need to consider null expectations in investigations of the geographical variation in body size together with the probable environmental mechanisms underlying spatial variation in average size. Moreover, it suggests that in the South African avifauna, spatial variation in the body sizes of assemblages may be determined indirectly by the factors that influence geographical variation in species richness.

During normal development, tadpoles of Xenopus laevis demonstrate large variations in body size that are carried through metamorphosis. This variation in size exists at the stages when lumbar lateral motor column (L-LMC) motoneurons are produced and when neuronal cell death in this neuron population occurs. Body size, hindlimb size, motoneuron number, and motoneuron size (i.e., neuron nuclear cross-sectional area) were measured in animals from three developmental stages: one prior to significant amounts of cell death, one at the peak rate of cell death, and one after cell death. The hypothesis that neuron population size is matched to peripheral size was tested by using the natural size variation found at each of these stages. The ranges of values for the measurements at the three stages were large. Significant correlations between body size and motoneuron number, as well as between motoneuron number and muscle fiber number, were present after cell death. Since these correlations emerged as cell death reduced neuron numbers, size matching may have occurred and cell death may have adjusted the L-LMC motoneuron population's size to variation in body size. In addition to the correlations between body size and motoneuron number at the end of cell death, neuron numbers before and after cell death were significantly correlated among groups of siblings. The possibility that the number of neurons after cell death was also influenced by differences in the number of L-LMC progenitors is discussed.

Intraspecific variability and species turnover drive variations in Collembola body size along a temperate-boreal elevation gradient

1. The egg size of insects can vary depending on maternal body size or resource status, and it may influence offspring body size by determining initial resource level.2. The giant rhinoceros beetle Trypoxylus dichotomus exhibits considerable variation in body size, some of which is attributed to the variation in larval food (humus) quality, although a substantial amount of variation in body size remains unexplained. In the present study, changes in the egg size and offspring body size in response to several maternal variables were examined (i.e. body size, age, and, nutritional status).3. Nutritional intake of the females during the adult stage did not affect the egg size. Larvae hatched from small eggs partially recovered from the initial disadvantage during their ontogenetic processes by increasing growth rate (i.e. compensatory growth); however, there was still a positive relationship between egg size and pupal body size.4. Older females produced small eggs, but because of compensatory growth, the pupae were no longer small. By contrast, due to a lack of compensatory growth, small females produced small eggs as well as small pupae.5. These results suggest that maternal body size affects offspring body size through effects on egg size. This transgenerational effect may account for some of the variation in adult body size of T. dichotomus.

Hypotheses that explain geographic variation in body size were examined using cranial measurements of 950 bobcats (Lynx rufus) from western North America. Bobcats were divided into 25 geographic localities of similar habitats and landform (based on ecoregions). Principal component analyses were used to derive a single estimate of size from scores on the first principal component. Males and females were examined separately, because they were significantly dimorphic in body size and because sex and locality exhibited a significant interaction. We expected that female body size would best reflect environmental influences, because male size may be influenced by sexual selection. We found significant geographic variation in bobcat body size, with about 44% of the variation in males and 47% of the variation in females accounted for by comparison among the localities. We also found that variation in body size was associated with Bergmann's rule, as indicated by significant multiple regression of body size of males (R2 = 0.426) and females (R2 = 0.480) on latitude and elevation. Using correlation and regression analyses, we examined the association of body size with selected environmental variables that represent the classical physiological explanation of Bergmann's rule, James' moisture-humidity modification of Bergmann's rule, Rosenzweig's productivity hypothesis, and Boyce's seasonality hypothesis. Only the productivity hypothesis was not supported. The relative strengths of associations suggested, however, that James' modification was better supported than the classical explanation for Bergmann's rule. Path analyses permitted further discrimination of hypotheses, and only the seasonality hypothesis received significant support. As expected, this support was only evident for females. Path analysis may provide a tool for evaluating relative strengths of competing but correlated explanations of geographic variation.

An increase in genetic variation in body size has often been observed under stress; an increase in dominance variance and interaction variance as well as in additive genetic variance has been reported. The increase in genetic variation must be caused by physiological mechanisms that are specific to adverse environments. A model is proposed to explain the occurrence of an increase in genetic variation in body size in Drosophila at extreme temperatures. The model has parameters specific to the low- and high-temperature regions of the viable range. Additive genetic variation in the boundary temperatures leads to a marked increase in additive genetic variation in development rate and body size at extreme temperatures. Additive genetic variation in the temperature sensitivity in the low- and high-temperature regions adds non-additive genetic variation. Development rate shows patterns in additive genetic variation that differ from the patterns of genetic variation in body size; therefore, the genetic correlation between development rate and body size changes sign repeatedly as a function of temperature. The existence of dominance in the genetic variation in the boundary temperatures or in the low- and high-temperature sensitivities leads to a higher total genetic variance due to higher dominance and interaction variance, for both development rate and body size.

Although variation in body size has been recently reported in stingless bees (Meliponini), empirical evidence evaluating possible factors related to such variation is lacking, and thus it is not clear if it may have an adaptive significance. We evaluated if variation in the body size and weight of workers of stingless bees fluctuates across a seasonal pattern and if this could be related to characteristics of the food consumed during the larval stage. The weight of larval provisions, their protein, and sugar content were evaluated in four colonies of Nannotrigona perilampoides every 2 months across 1 year. Worker-destined larvae from the same combs were allowed to develop and were sampled as callow workers to determine their weight and size using morphometric data. The weight and size of workers were highly correlated and varied across the seasons in established colonies, suggesting that size variation cycles across the year in stingless bees. An increase in the protein content and, to a lesser degree, the quantity of larval food were positively linked to variation in body weight and size; food with richer protein content resulted in larger and heavier workers. This study provides the first evidence of an effect of the quantity and composition of larval food on the size of workers in stingless bees. Although body weight and size of workers differed across seasons, they were not readily noticeable as changes seem to occur as a continuum across the year. Since size polymorphism was of a larger magnitude across time but not within age cohorts and as it was highly determined by food resources, it may not be an adaptive feature in stingless bees. However, more studies are needed to determine the role of the cyclical change in worker body size on colony performance and thus its adaptive significance in stingless bees.

Intra-specific body size variation is common and often is assumed to be adaptive. Studies of body size variation among sites should include or consider environmental and ecological variables in their designs. Additionally, reciprocal transplant or common garden studies will support which variables are really contributing to the observed body size variation. This study analyzed the microgeographic body size variation in Anolis mariarum, a small lizard endemic to Antioquia, Colombia. Parameters such as body size, shape, and lepidosis variation were quantified in 217 adult A. mariarum, belonging to six populations separated by less than 80km. Results showed that significant body size variation was not related to differences among sites in mean annual temperature, but covaried with mean annual precipitation, with the largest individuals occurring in dryer sites. Mark-recapture data obtained from 115 individuals from both the wettest and dryest sites from October 2004 to April 2005 showed that growth rates were higher at the latter. Eight males from each site were captured at the end of the mark-recapture study and reared for two months under identical conditions in a common garden study. Individuals from both sites grew faster when reared in the laboratory with food provided ad libitum. Although growth rates of males from the two populations did not differ significantly in the laboratory, males from the dryest site still maintained a significantly larger asymptotic body size in their growth trajectories. Multivariate analyses also demonstrated that both males and females from the six populations differed in terms of body shape and lepidosis. However, only female body size was found to covary significantly with an environmental gradient (precipitation). A. mariarum does not conform to Bergmann's rule, but the relationship found between mean body size and asympotic growth with mean annual precipitation at these sites needs further analysis. Generally, studies of intra-specific body size variation should consider a number of additional phenotypic traits to provide stronger baseline information on the degree of overall divergence among populations, including those likely to be selectively neutral, before interpreting results of analyses on the body size differences.

Many animal species show considerable geographic variation in average adult body sizes. Although this phenomenon offers a unique opportunity to tease apart the factors influencing body size, interpretation has proved to be difficult because several processes may contribute to the observed variation (e.g., Andrews, 1976; Case, 1978; Dunham et al., 1978). First, average body size is influenced by adult survival rates in species in which growth continues after maturity: animals may be smaller in one area than another simply because they are, on average, younger (e.g., King, 1989). Second, even if survivorships are similar among areas, so that the body-size differences are due to differences in growth patterns, there are two possible explanations for such variation in growth. Body sizes may differ either because of local genetic modifications (possibly due to adaptation) or because of a direct phenotypic effect of differing food availability on growth rates (e.g., Berry et al., 1987; Dobson and Murie, 1987; Ebenhard, 1 990). To distinguish among these alternative interpretations, we need two kinds of data on growth trajectories of animals: first from the field (to determine whether the populations differ in actual growth patterns, rather than simply survivorships) and second the response to experimental manipulation of food supply (to determine whether the observed differences in growth rates are due to genetic differences or phenotypic plasticity). This paper presents information on European grass snakes. We show that body sizes and the degree of sexual difference in body size are greatly reduced in an island population compared to the nearby mainland, that these differences are due to modified growth patterns and not just survivorship, and that the low growth rates and small asymptotic body sizes of the island snakes are a phenotypic response to local conditions (probably, low food availability). Insular populations of snakes offer some of the most dramatic examples of geographic variation in body size (e.g., Case, 1978; Schwaner, 1985; King, 1989; Shine, 1987; Schwaner and Sarre, 1988; Hasegawa and Moriguchi, 1989; Forsman, 1991). For example, Schwaner (1985) showed that body masses of adult Australian tigersnakes varied up to tenfold among adjacent islands. Correlational analyses suggest that predators attain larger sizes in areas where larger species of prey are available, and this may be true both for snakes (Schwaner, 1985; Hasegawa and Moriguchi, 1989) and for mammalian predators (Gittleman, 1985; Erlinge, 1987). Experimental studies on mammals have shown that geographic variation in body sizes may be due both to phenotypic plasticity (e.g., Dobson and Murie, 1987) and to local adaptation (e.g., Berry et al., 1987; Ebenhard, 1990). The only experimental study to address the determinants of such differences in snakes has been that of Barnett and Schwaner (1984), who raised juvenile tigersnakes. These authors documented rapid growth in captive snakes from a giant population, but obtained no comparable information on snakes from dwarf populations. We studied two populations of a nonvenomous natricine colubrid species, the grass snake (Natrix natrix), which is abundant over much of Europe (Arnold and Burton, 1978). The mainland study area was near Maryd, 15 km south of Lund in southern Sweden (55?40'N, 13?30'E). The area contains a mixture of arable land, grazed meadow, and mixed deciduous forest. Detailed data have already been published on body sizes, sexual size dimorphism, growth rates, diets and reproductive biology of grass snakes from this area (Madsen, 1983, 1987). Those papers also describe the methods used to capture, mark and measure snakes, and to obtain prey items by forced regurgitation. The same methods were used for the study of island snakes. Our island population was on Hallands Vadero (56?27'N, 12?44'E), a small (2.6 kM2) island approximately 3 km from the Swedish coast. One quarter of the island is forested, with the remainder consisting of meadows bordered by blackthorn, stony areas with juniper, and bare rock (Madsen and Stille, 1988). The two study areas are approximately 100 km apart. Geological evidence suggests that the two areas probably have been separated for several thousand years, since glaciation-induced reductions in sea level (Devoy, 1987). However, it is possible that the effective period of separation may have been longer than this (e.g., these oviparous snakes may not have existed in this region during glacial periods) or considerably briefer (due to fortuitous dispersal of snakes from one area to another). Male snakes from the island were similar to the mainland animals in mean SVL (Fig. 1; N = 22, 41; means = 52.2 versus 53.0 cm, t = 0.40, df= 61, P = 0.69) but females were much smaller on the island (N = 28, 44; means 59.7 versus 69.9 cm, t = 4.14, df= 70, P < 0.001). Indeed, some male snakes from the mainland actually attained larger sizes than did any of the females from the island population, although the mean values were lower (Fig. 1). Two-factor analysis

Microhabitat characteristics are expected to influence the distribution of stream fish species at fine spatial scales (e.g., within riffle segments). Body size is probably the most important trait that constrains microhabitat occupation by fish, but the effect of intraspecific variation has been understudied. We investigated how physical microhabitat characteristics affect species and body size distribution of fish within a stream riffle segment in a coastal subtropical drainage of Brazil. Fishes were sampled by electrofishing 56 riffle plots along a 730-m long stream segment. Species composition was significantly related to four microhabitat characteristics: substrate size, flow velocity, distance to margin and depth. In addition, mean body size increased with increasing substrate size and depth of microhabitat sampling plots. However, when including species identity in linear mixed-effects models (LMM), we observed a different relationship between body size and microhabitat characteristics, but most of the variation was explained by species identity. Thus, we fitted LMMs separately for each species and found species-specific relations between intraspecific variation in body size and microhabitat characteristics. The low variation explained in the models suggests that other fine scale factors, such as biotic interactions and dispersal from adjacent habitat patches, should be incorporated in modeling microhabitat use by stream fish. Our findings suggest that body size is important by itself, but intraspecific variation in body size also constrains microhabitat use differently for each species, which may depend on other species-specific traits, such as morphology, behavior and life history.

The body size of an adult insect is strongly determined by the environmental factors to which it is exposed during growth and development. Insect species confronted with a high environmental variability across their geographical range (i.e. wide ecological niche breadth) may therefore reveal broader variation in body size than those species which are more specialised (i.e. narrow ecological niche). In this study, we aim to investigate whether characteristics related to the ecological niche breadth of a holometabolous insect species (i.e. its ecological specialisation) affect its intraspecific variation in adult body size. By using European geometrid moths as a model group, we specifically tested whether latitudinal range size, larval resource use and voltinism affect intraspecific body size variation. We hypothesised that body size variation will increase along with latitudinal range and larval diet breadth. We further expected that univoltine species reveal a lower body size variation compared to those with multiple generations per year. To test these hypotheses, we compiled a comprehensive trait database for 631 species of European geometrid moths from literature, including information on adult body size, life history and distribution. We further reconstructed a molecular phylogeny including all analysed geometrid species and applied phylogenetic comparative methods in order to test our predictions. In support of our hypotheses, we found that intraspecific size variation is positively related to latitudinal range size and larval diet breadth, and that multivoltine species reveal a higher heterogeneity in body size than taxa with a strictly univoltine life style. Based on our results, we demonstrated that intraspecific body size variation in geometrid moths is negatively related to ecological specialisation. We further suggest that increased variation in body size with increasing niche breadth is a general pattern, which likely applies to many other insect groups as well. This assumption, however, demands further empirical scrutiny.

Most ecological studies attempting to understand causes of population dynamics and community structure disregard intraspecific trait variation. We quantified the importance of natural intra-cohort variation in body size and density of juveniles for recruitment of a sessile marine organism, the barnacle Semibalanus balanoides. Barnacles are representative of species organised in metapopulations, that is, as open local populations connected by larval dispersal. We tracked the individual growth and survival of a cohort of juvenile barnacles from two shores of North Wales. Barnacles settled as larvae in spring of 2002 on previously cleared rock. The density of these new recruits was experimentally manipulated in June and randomly selected individuals were monitored from June to October to evaluate the role of barnacle size and density in predicting survival. In doing so we characterised density at three spatial scales (quadrat: 25 cm2, cells within quadrats: 25 mm2 and neighbourhood: number of neighbours in physical contact with the target barnacle). At all scales, variations in juvenile body size exacerbated the effect of density-dependent mortality on population size. While density-dependent mortality was very intense in the small-sized individuals, large-sized individuals experienced very weak density-dependent mortality and showed high survival rates. Using the concept of 'Jensen inequality', we show that important biases in estimations of survival, based on population size only, occur at high barnacle densities, where survival is low. Our study highlights the role of body size variation in understanding dynamics of open populations.

Patterns of abundance across space and time, and intraspecific variation in body size, are two species attributes known to influence diet breadth and the structure of interaction networks. Yet, the relative influence of these attributes on diet breadth is often assumed to be equal among taxonomic groups, and the relationship between intraspecific variation in body size on interaction patterns is frequently neglected. We observed bee–flower interactions in multiple locations across Montana, USA, for two growing seasons and measured spatial and temporal patterns of abundance, along with interspecific and intraspecific variation in body size for prevalent species. We predicted that the association between spatial and temporal patterns of abundance and intraspecific variation in body size, and diet breadth, would be stronger for bumble bee compared to non-bumble bee species, because species with flexible diets and long activity periods can interact with more food items. Bumble bees had higher local abundance, occurred in many local communities, more intraspecific variation in body size, and longer phenophases compared to non-bumble bee species, but only local abundance and phenophase duration had a stronger positive association with the diet breadth of bumble bee compared to non-bumble bee species. Communities with a higher proportion of bumble bees also had higher intraspecific variation in body size at the network-level, and network-level intraspecific variation in body size was positively correlated with diet generalization. Our findings highlight that the association between species attributes and diet breadth changes depending on the taxonomic group, with implications for the structure of interaction networks.

Context Seasonal migration and movements of bats have important implications for their conservation. The southern bent-winged bat (Miniopterus orianae bassanii), a critically endangered cave-dwelling taxon in Australia, has been described as undertaking regional-scale migration between maternity and non-breeding caves. Aims To describe the seasonal cycle of movements by the southern bent-winged bat, including migration and congregation events of different sex- and age-classes in the population. Methods We tagged a total of 2966 southern bent-winged bats with passive integrated transponder (PIT) tags. Antennas were used to detect bats in flight at a major maternity cave and a key non-breeding cave in south-east South Australia, from January 2016 to August 2019. We used capture–resight histories to visualise population patterns and model the daily encounter probability for each sex- and age-class at the respective roost sites. Key results Bats congregated at the maternity cave for most of the year, with different seasonal patterns among sex- and age-classes. Seasonal movements were associated with behaviour over winter months: most of the population dispersed from the maternity cave from May and a staged return occurred among population classes from July through September. A previously undescribed movement occurred in adult females and juveniles each year: these classes left the maternity cave in late summer, when juveniles became independent, and returned in early mid-autumn, later undertaking winter dispersal. Complex underlying movements of individuals occurred throughout the year, with individuals able to fly 72 km between roosting caves in just a few hours. Conclusions Seasonal movements are a key aspect of the life history of this taxon. The newly reported movement of adult females and juveniles conforms to the maternal guidance hypothesis, whereby mothers guide their young to suitable non-breeding caves and hibernation sites. In addition to seasonal movements, some individuals moved 72 km between caves multiple times over short time periods, including on successive nights. This 72-km overnight flight distance more than doubles the previous distance used to inform management buffer zones. Extended congregation of bats at the maternity cave highlights resource limitation in the surrounding area as a potential threat to this population. Implications The dynamic nature of the population has implications for the management of emerging risks, including mortality at windfarms and potential rapid spread of the exotic white-nose syndrome.