Ecological correlates of range shifts of Late Pleistocene mammals

Understanding and predicting how species' distributions will shift as climate changes are central questions in ecology today. The late Quaternary of North America represents a natural experiment in which we can evaluate how species responded during the expansion and contraction of the glaciers. Here, we ask whether species' range shifts differ because of taxonomic affinity, life-history traits, body size or topographic heterogeneity and whether the species survived the megafaunal extinction. There was no difference in range shifts between victims and survivors of the megafaunal extinction. In general, the change in the size of a species' range is not well correlated with any of the ecological or life-history traits evaluated. However, there are significant relationships between some variables and the movements of the centroids of ranges. Differences in the distances shifted exist among orders, although this is probably a result of body size differences as larger bodied species show larger shifts. Although there are a few exceptions, the distance that species shifted their range was weakly correlated with life-history traits. Finally, species in more topographically heterogeneous areas show smaller shifts than species in less-diverse areas. Overall, these results indicate that when trying to predict species range shifts in the future, body size, lifespan and the topographic relief of the landscape should be taken into account.     

Comparative analysis of feeding habits and dietary niche breadth in skates: the importance of body size,snout length,and depth

Skates (Elasmobranchii, Rajiformes) are a morphologically conservative group of bentophagous chondrichthyan fishes with a high degree of endemism, that occur on marine soft bottoms. Subtle morphological aspects and bathymetric distribution are traits that vary among skate species that could have implications for their feeding ecology. We test how body size, snout length and bathymetric distribution influence the feeding habits and dietary niche breadth in skates using data on 71 species taken from the literature. We hypothesized that snout length has an effect on diet composition. We also hypothesized that dietary niche breadth increases with increasing depth range and decreases with increasing body size of skate species. Generalized additive models for location scale and shape were fitted with taxonomic level (genera nested within family) included as a random effect term in each model. A model selection approach to test the level of support for alternative models was applied. We found that skate species that forage on large prey have the largest body size and skate species with the smallest body size prey on small and medium-sized invertebrates. The results indicated that body size has an effect on feeding habits of skates, whereas an effect of snout length was not supported. Bathymetric variables have an effect on the diet of skates. Our prediction that dietary niche breadth increases with increasing depth range and decreases with increasing body size of skate species was supported in part: in a first phase the relationship between dietary niche breadth and body size is positive, then in a second phase, including species larger than 1000 mm total length, the relationship become negative.     

Ecological and evolutionary drivers of range size in Coenagrion damselflies

Geographic range size is a key ecological and evolutionary characteristic of a species, yet the causal basis of variation in range size among species remains largely unresolved. One major reason for this is that several ecological and evolutionary traits may jointly shape species' differences in range size. We here present an integrated study of the contribution of ecological (dispersal capacity, body size and latitudinal position) and macroevolutionary (species' age) traits in shaping variation in species' range size in Coenagrion damselflies. We reconstructed the phylogenetic tree of this genus to account for evolutionary history when assessing the contribution of the ecological traits and to evaluate the role of the macroevolutionary trait (species' age). The genus invaded the Nearctic twice independently from the Palearctic, yet this was not associated with the evolution of larger range sizes or dispersal capacity. Body size and species' age did not explain variation in range size. There is higher flight ability (as measured by wing aspect ratio) at higher latitudes. Species with a larger wing aspect ratio had a larger range size, also after correcting for phylogeny, suggesting a role for dispersal capacity in shaping the species' ranges. More northern species had a larger species' range, consistent with Rapoport's rule, possibly related to niche width. Our results underscore the importance of integrating macroecology and macroevolution when explaining range size variation among species.     

Larger Body Size at Metamorphosis Enhances Survival,Growth and Performance of Young Cane Toads (Rhinella marina)

Body size at metamorphosis is a key trait in species (such as many anurans) with biphasic life-histories. Experimental studies have shown that metamorph size is highly plastic, depending upon larval density and environmental conditions (e.g. temperature, food supply, water quality, chemical cues from conspecifics, predators and competitors). To test the hypothesis that this developmental plasticity is adaptive, or to determine if inducing plasticity can be used to control an invasive species, we need to know whether or not a metamorphosing anuran’s body size influences its subsequent viability. For logistical reasons, there are few data on this topic under field conditions. We studied cane toads (Rhinella marina) within their invasive Australian range. Metamorph body size is highly plastic in this species, and our laboratory studies showed that larger metamorphs had better locomotor performance (both on land and in the water), and were more adept at catching and consuming prey. In mark-recapture trials in outdoor enclosures, larger body size enhanced metamorph survival and growth rate under some seasonal conditions. Larger metamorphs maintained their size advantage over smaller siblings for at least a month. Our data support the critical but rarely-tested assumption that all else being equal, larger body size at metamorphosis is likely to enhance an individual’s long term viability. Thus, manipulations to reduce body size at metamorphosis in cane toads may help to reduce the ecological impact of this invasive species.     

Sexual size dimorphism in musteloids: An anomalous allometric pattern is explained by feeding ecology

Rensch's rule states that sexual size dimorphism (SSD) increases with body size in taxa where males are larger, and decreases when females are larger. The dominant explanation for the trend is currently that competitive advantage for males is greater in larger individuals, whereas female size is constrained by the energetics of rearing offspring. This rule holds for a variety of vertebrate taxa, and opposing trends are rare. We examine the allometry of SSD within the Musteloidea and demonstrate a hypo‐allometry contrary to Rensch's rule, with lower SSD associated with larger body size. We provide evidence that feeding ecology is involved. Where diet promotes group‐living, the optimal strategy for the males of larger species is often not to attempt to defend access to multiple females, obviating any competitive advantage of relatively greater size. We conclude that the effect of feeding ecology on mating systems may be a hitherto neglected factor explaining variation in SSD.     

Pattern and process in the geographical ranges of freshwater fishes

North American freshwater fishes were studied to determine whether they displayed the same relationships between log (geographical range size) and log (body size) and the same pattern of range shape as found among North American birds and mammals. The forces that produce these patterns were also investigated. The log (geographical range size) : log (body size) relationship was analysed for 121 North American freshwater fish species. Thirty‐two imperilled species were compared with 89 non‐imperilled species to determine if the overall relationship could result from differential extinction. Range geometries were analysed, within and among habitat guilds, to determine if general patterns could be detected. The log (geographical range size) : log (body size) pattern among freshwater fish species was triangular and qualitatively similar to that found for North American birds and mammals. The results suggest that below a minimum geographical range, the likelihood of extinction increases dramatically for freshwater fishes and that this minimum range size increases with body size. The pattern of fish species’ range shapes differs from that found for other North American vertebrate taxa because, on average, fish possess much smaller ranges than terrestrial species and most fish species’ geographical ranges extend further on a north–south axis than on an east–west axis. The log (geographical range size) : log (body size) pattern reveals that fish species’ geographical ranges are more constrained than those of terrestrial species. The triangular relationship may be caused by differential extinction of species with large bodies and small geographical ranges as well as higher speciation rates of small‐bodied fish. The restricted geographical ranges of freshwater fishes gives them much in common with terrestrial species on oceanic islands. Range shape patterns within habitat guilds reflect guild‐specific historical and current ecological forces. The overall pattern of range shapes emerges from the combination of ecologically different subunits.     

Species distribution,ecology, abundance,body size and phylogeny originate interrelated rarity patterns at regional scale

The most pervasive macroecological patterns concern (1) the frequency distribution of range size, (2) the relationship between range size and species abundance and (3) the effect of body size on range size. We investigated these patterns at a regional scale using the tenebrionid beetles of Latium (Central Italy). For this, we calculated geographical range size (no. of 10‐km square cells), ecological tolerance (no. of phytoclimatic units) and abundance (no. of sampled individuals) using a large database containing 3561 georeferenced records for 84 native species. For each species, we also calculated body mass and its ‘phylogenetic diversity’ on the basis of cladistic relationships. Frequency distribution of range size followed a log‐normal distribution as found in many other animal groups. However, a log‐normal distribution accommodated well the frequency distribution of ecological tolerance, a so far unexplored issue. Range size was correlated with abundance and ecological tolerance, thus supporting the hypothesis that a positive correlation between distribution and abundance is a reflection of interspecific differences in ecological specialization. Larger species tended to have larger ranges and broader ecological tolerance. However, contrary to what known in most vertebrates, not only small‐sized, but also many medium‐to‐large‐sized species exhibited great variability in their range size, probably because tenebrionids are not so strictly influenced by body size constraints (e.g. home ranges) as vertebrates. Moreover, in contrast to other animals, tenebrionid body size does not influence species abundances, probably because these detritivorous animals are not strongly regulated by competition. Finally, contrary to the assumption that rare species should be mainly found among lineages that split from basal nodes, rarity of a tenebrionid species was not influenced by the phylogenetic position of its tribe. However, lineages that split from more basal nodes had lower variability in terms of species geographical distribution, ecological tolerance and abundance, which suggests that lineages that split from more basal nodes are not only morphologically conservative but also tend to have an ecological ‘inertia’.     

Ecological specialization in fossil mammals explains Cope's rule

Cope's rule is the trend toward increasing body size in a lineage over geological time. The rule has been explained either as passive diffusion away from a small initial body size or as an active trend upheld by the ecological and evolutionary advantages that large body size confers. An explicit and phylogenetically informed analysis of body size evolution in Cenozoic mammals shows that body size increases significantly in most inclusive clades. This increase occurs through temporal substitution of incumbent species by larger-sized close relatives within the clades. These late-appearing species have smaller spatial and temporal ranges and are rarer than the incumbents they replace, traits that are typical of ecological specialists. Cope's rule, accordingly, appears to derive mainly from increasing ecological specialization and clade-level niche expansion rather than from active selection for larger size. However, overlain on a net trend toward average size increase, significant pulses in origination of large-sized species are concentrated in periods of global cooling. These pulses plausibly record direct selection for larger body size according to Bergmann's rule, which thus appears to be independent of but concomitant with Cope's.     

Variability and size in mammals and birds

Body size, its variability, and their ecological correlates have long been important topics in evolutionary biology. Yet, the question of whether there is a general relationship between size and size-relative variability has not previously been addressed. Through an analysis of body-mass and length measurements from 65 074 individuals from 351 mammalian species, we show that size-relative variability increases significantly with mean species body size. Analysis of mean body mass and standard deviations for 237 species of birds revealed the same pattern. We present three plausible alternatives explanations and eliminate several others. Of these, the hypothesis that the increase in size-relative variability with mean body mass is related to the scaling of body mass components is most strongly supported. In effect, larger mammals and birds are more variable because their body mass is composed to greater relative degree of components with higher intrinsic variability (bone, fat, and muscle). In contrast, smaller mammals and birds have lower body mass variability because they are composed to a greater relative extent of components (viscera and nervous system) in which size variation is more highly constrained by energetic and functional factors.     

Breeding ecology and oviposition site selection of black-spotted pond frogs (Rana nigromaculata) in Ningbo, China

The black-spotted pond frog (Rana nigromacu-lata) is one of the most widely distributed species in China. However, there have been only a few qualitative descrip-tions of their breeding ecology and oviposition site selec-tion. From 2004 to 2006, we investigated the breeding ecology and oviposition site selection of K nigromaculata in Ningbo, China, quantitatively. Analyses of breeding ecology show that: (1) mean frog density in the breeding season was 0.0903 ± 0.0029 individuals per meter (n = 11) (mean ± SE); (2) R. nigromaculata was a sexually dimorphic species, with females significantly larger than males in both body weight and snout-vent length; (3) the clutch size averaged 4643.04± 235.96 eggs (range 1546-7897, n = 50); and (4) the egg size ranged from 1.50 to 1.74 mm in diameter, with an average egg size of 1.6050 ± 0.0046 mm (n = 226). Oviposition sites differed significantly from random sites in percentages of water, bare ground and vegetation cover, water depth (cm), water temperature (°C) and water turbidity. Rana nigro-maculata preferred microhabitats with higher percentages of water and vegetation cover, while it avoided microha-bitats with deeper water. The results suggest that micro-habitats with higher percentages of water and vegetation cover, but not deeper water, should be priorities for pro-tection and conservation of the breeding habitats of R. nigromaculata.     

HAVE MALE AND FEMALE GENITALIA COEVOLVED? A PHYLOGENETIC ANALYSIS OF GENITALIC MORPHOLOGY AND SEXUAL SIZE DIMORPHISM IN WEB‐BUILDING SPIDERS (ARANEAE: ARANEOIDEA)

Sexual size dimorphism (SSD) can strongly influence the evolution of reproductive strategies and life history. If SSD is extreme, and other characters (e.g., genitalic size) also increase with size, then functional conflicts may arise between the sexes. Spiders offer an excellent opportunity to investigate this issue because of their wide range of SSD. By using modern phylogenetic methods with 16 species of orb-weaving spiders, we provide strong evidence for the "positive genitalic divergence" model, implying that sexual genitalic dimorphism (SGD) increases as SSD increases. This pattern is supported by an evolutionary mismatch between the absolute sizes of male and female genitalia across species. Indeed, our findings reveal a dramatic reversal from male genitalia that are up to 87x larger than female genitalia in size-monomorphic species to female genitalia that are up to 2.8x larger in extremely size-dimorphic species. We infer that divergence in SGD could limit SSD both in spiders, and potentially in other taxa as well. Further, male and female body size, as well as male and female genitalia size, are decoupled evolutionarily. Finally, we show a negative scaling (hypoallometry) of male and female genitalic morphology within sexes. Evolutionary forces specific to each sex, such as larger female size (increased fecundity) or smaller male size (enhanced mate-searching ability), may be balanced by stabilizing selection on relative genitalic size.     

Range size in North American Enallagma damselflies correlates with wing size

1. Cross-species macroecological comparisons in freshwater invertebrates have been restricted by a lack of large-scale distributional data, and robust phylogenies. Here, we use data from the OdonataCentral database to explore body length–range size and wing length–range size relationships in damselflies from the genus Enallagma ; the recent publication of a phylogeny for this group meant that, as well as a cross-species analysis, we were able to assess relationships in a phylogenetically controlled manner.
2. For cross-species comparisons, only wing length showed significant (positive) regression relationships with range size and occupancy, although the inclusion of body length in multiple regressions increased the fit of the models. Damselflies with larger wings relative to their body length had larger distributions, a result confirmed by a significant positive relationship between range size and residuals from the regression of wing size on body size.
3. For the phylogenetically controlled analyses, only wing length contrast scores were significantly related to distribution patterns and entered into regression models; the significant positive relationships between wing length contrasts and both range size and occupancy contrasts suggested that evolutionary increases in wing length had occurred alongside range expansions.
4. Together these results suggest that species of Enallagma with larger wings (both absolute and relative to body length) tend to be more widely distributed in North America and that the evolution of wing size may have played a role in range expansion. No such relationships were evident for body size. We discuss the potential importance of wing morphometrics for studying the evolutionary ecology of freshwater insects.     

Geographical and ecological patterns of range size in North American trees

We analysed the degree to which spatial, ecological and phylogenetic factors influenced geographic gradients in range size for all trees native to North America. We compared observed range sizes, measured in either one (latitudinal or longitudinal extent) or two dimensions (range area), with three null models that respected constraints imposed by the geometry of the continent; we tested whether climatic gradients could account for increasing range size with latitude – a Rapoport effect – in North American trees; and whether variation in range size was associated with phylogeny or ecological characteristics of species. We documented an excess of species with small ranges and a lack of widely distributed species compared with null expectations both with and without environmental constraints. We found evidence for a Rapoport effect in North American trees, at different taxonomic levels and for different groups defined by ecological characteristics. This pattern is not an outcome constrained by continental shape and does not depend on the metric for range size, but it is consistent with the climatic variability hypothesis proposed to explain the Rapoport pattern. Finally we showed that trees east of the Rocky Mountains have larger ranges on average than trees to the west or in Mexico and that the proportion of deciduous compared to evergreen species increases with range size. Variation in range size among North American trees is not spatially random, and has a phylogenetic and ecological signal. Consistent with the climatic variability hypothesis, range size of North American trees increases with increasing variability in annual temperature range at higher latitudes.     

宏生态学(Macroecology)及其研究

宏生态学是生态学与其他宏观学科不断交叉和融合后的产物。它以个体、种群和物种的生态特征在大时空尺度上的格局和变化规律为主要研究内容,它比其他生态学更强调归纳和推论,也更依赖数据的积累。近年来,宏生态学在对物种一面积关系进行探讨的基础上,对生物类群间的物种数量的协同变化以及物种和高级分类单元间的关系等进行了新的研究;宏生态学试图将有机论和个体论结合来探讨和总结群落结构中的物种组成规律;并对物种多度和分布格局间的关系从生态位和异质种群角度进行新的解释;个体大小频次分布规律是宏生态学一重要内容,对其深入研究和探讨已与物种多度、能量、分布面积、历史起源等多方面特征相结合,并得到一些普遍性规律;最后,宏生态学还探讨物种在地理区域上的普遍性的分布模式,并对其假说进行检验和探讨。宏生态学在中国还处于刚起步阶段,但中国具有资源的优势,并具有一定的数据积累,将在宏生态学研究中发挥越来越重要的作用。     

What ecological factors shape species-area curves in neutral models?

Understanding factors that shape biodiversity and species coexistence across scales is of utmost importance in ecology, both theoretically and for conservation policies. Species-area relationships (SARs), measuring how the number of observed species increases upon enlarging the sampled area, constitute a convenient tool for quantifying the spatial structure of biodiversity. While general features of species-area curves are quite universal across ecosystems, some quantitative aspects can change significantly. Several attempts have been made to link these variations to ecological forces. Within the framework of spatially explicit neutral models, here we scrutinize the effect of varying the local population size (i.e. the number of individuals per site) and the level of habitat saturation (allowing for empty sites). We conclude that species-area curves become shallower when the local population size increases, while habitat saturation, unless strongly violated, plays a marginal role. Our findings provide a plausible explanation of why SARs for microorganisms are flatter than those for larger organisms.     

The 'island rule' in birds: medium body size and its ecological explanation

Do birds show a different pattern of insular evolution from mammals? Mammals follow the ''island rule'', with large-bodied species getting smaller on islands and small-bodied species getting bigger. By contrast, the traditional view on birds is that they follow no general island rule for body size, but that there is an insular trend for large bills. Insular shifts in feeding ecology are, therefore, widely assumed to be the primary cause of divergence in island birds. We use a comparative approach to test these ideas. Contrary to the traditional view, we find no evidence for increased bill size in insular populations. Instead, changes in both bill size and body size obey the ''island rule''. The differences between our results and the traditional view arise because previous analyses were based largely on passerines. We also investigate some ecological factors that are thought to influence island evolution. As predicted by the traditional view, shifts in bill size are associated with feeding ecology. By contrast, shifts in body size are associated with the potential for intraspecific competition and thermal ecology. All these results remain qualitatively unchanged when we use different methods to score the ecological factors and restrict our analyses to taxa showing pronounced morphological divergence. Because of strong covariation between ecological factors, however, we cannot estimate the relative importance of each ecological factor. Overall, our results show that the island rule is valid for both body size and bill length in birds and that, in addition to feeding ecology, insular shifts in the level of intraspecific competition and the abiotic environment also have a role.     

Species traits predict assembly of mayfly and stonefly communities along pH gradients

Much recent ecological research has centred on the interrelations between species diversity and ecological processes. In the present study, I show how species traits may aid in comprehending ecology by studying the link between an environmental variable and functional traits. I examined the composition of species traits with a theoretically underpinned relationship to ecological processes along a pH gradient. I focused on body size, reproductive output, life cycle length and feeding habit of mayflies and stoneflies. In mayfly assemblages, I found smaller body size, greater reproductive output, faster life cycles and a larger proportion of gathering collectors and scrapers with increasing pH. In stonefly assemblages, I found smaller body size, greater reproductive output and faster life cycles at sites with a history of long-term natural acidification, but no clear trends in feeding habits and in most traits where acidification is anthropogenic. The results suggest that mayflies and stoneflies exhibit different ecological functions following different ecological strategies. Mayflies follow an opportunistic strategy relative to stoneflies, likely facilitating high rates of ecological processes with respect to the autotrophic resource base at neutral sites. Relative to mayflies, stoneflies follow an equilibrium strategy contributing to ecological functioning in heterotrophic ecosystems and likely maintaining heterotrophic processes despite the erosion of species diversity in response to acidification. The rules governing an ecological community may be more readily revealed by studying the distribution of species traits instead of species diversity; by studying traits, we are likely to improve our understanding of the workings of ecological communities.     

The influence of sex and body size on food habits of a giant tropical snake, Python reticulatus

1. In many animal species, dietary habits shift with body size, and differ between the sexes. However, the intraspecific range of body sizes is usually low, making it difficult to quantify size-associated trophic shifts, or to determine the degree to which sex differences in diet are due to body-size differences. Large snakes are ideal for such a study, because they provide a vast range of body sizes within a single population.
2. More than 1000 Reticulated Pythons ( Python reticulatus ) from southern Sumatra were examined, with specimens from 1·5 to > 6 m in snout–vent length, and from 1 to 75 kg in mass. Females attained much larger body sizes than did conspecific males (maxima of 20 vs 75 kg, 5 vs 7 m), but had similar head lengths at the same body lengths.
3. Prey sizes, feeding frequencies and numbers of stomach parasites (ascarid nematodes) increased with body size in both sexes, and dietary composition changed ontogenetically. Small snakes fed mostly on rats, but shifted to larger mammalian taxa (e.g. pangolins, porcupines, monkeys, wild pigs, mouse deer) at 3–4-m body length.
4. Adult males and females showed strong ecological divergence. For some traits, this divergence was entirely caused by the strong allometry (combined with sexual size dimorphism), but in other cases (e.g. feeding frequency, dietary composition), the sexes followed different allometric trajectories. For example, females shifted from rats to larger mammals at a smaller body size than did conspecific males, and feeding frequencies increased more rapidly with body size in females than in males. These allometric divergences enhanced the degree of sex difference in trophic ecology induced by sexual size dimorphism.     

Contrasting selection pressure on body and weapon size in a polygynous megaherbivore

Sexual size dimorphism (SSD) is a common morphological trait in ungulates, with polygyny considered the leading driver of larger male body mass and weapon size. However, not all polygynous species exhibit SSD, while molecular evidence has revealed a more complex relationship between paternity and mating system than originally predicted. SSD is, therefore, likely to be shaped by a range of social, ecological and physiological factors. We present the first definitive analysis of SSD in the common hippopotamus (Hippopotamus amphibius) using a unique morphological dataset collected from 2994 aged individuals. The results confirm that hippos exhibit SSD, but the mean body mass differed by only 5% between the sexes, which is rather limited compared with many other polygynous ungulates. However, jaw and canine mass are significantly greater in males than females (44% and 81% heavier, respectively), highlighting the considerable selection pressure for acquiring larger weapons. A predominantly aquatic lifestyle coupled with the physiological limitations of their foregut fermenting morphology likely restricts body size differences between the sexes. Indeed, hippos appear to be a rare example among ungulates whereby sexual selection favours increased weapon size over body mass, underlining the important role that species-specific ecology and physiology have in shaping SSD.     

Life on the edge: carnivore body size variation is all over the place

Evolutionary biologists have long been fascinated by both the ways in which species respond to ecological conditions at the edges of their geographic ranges and the way that species'' body sizes evolve across their ranges. Surprisingly, though, the relationship between these two phenomena is rarely studied. Here, we examine whether carnivore body size changes from the interior of their geographic range towards the range edges. We find that within species, body size often varies strongly with distance from the range edge. However, there is no general tendency across species for size to be either larger or smaller towards the edge. There is some evidence that the smallest guild members increase in size towards their range edges, but results for the largest guild members are equivocal. Whether individuals vary in relation to the distance from the range edges often depends on the way edge and interior are defined. Neither geographic range size nor absolute body size influences the tendency of size to vary with distance from the range edge. Therefore, we suggest that the frequent significant association between body size and the position of individuals along the edge-core continuum reflects the prevalence of geographic size variation and that the distance to range edge per se does not influence size evolution in a consistent way.