On size and area: Patterns of mammalian body size extremes across landmasses

We describe a biogeographic pattern in which mammalian body size extremes scale with landmass area. The relationship between the largest and the smallest mammal species found on different landbridge islands, mountaintops and continents shows that the size of the largest species increases, while that of the smallest species decreases, with increase in the area of the landmass. We offer two possible explanations: (1) that the pattern is the result of sampling artefacts, which we call the ‘statistical artefact hypothesis’, or (2) that the pattern is the result of processes related to the way body size affects the number of individuals that a particular species can pack in a given area, which we call the ‘area-scaling hypothesis’. Our results point out that the pattern is not a statistical artefact resulting from random sampling, but can be explained by considering the scaling of individual space requirements and its effect on population survival on landmasses of different area. This revised version was published online in July 2006 with corrections to the Cover Date.     

Global change and carnivore body size: data are stasis

Aim  Global warming and other anthropogenic changes to the environment affect many aspects of biology and have often been invoked as causing body size changes in vertebrates. Here we examine a diverse set of carnivore populations in search of patterns in body size change that could reflect global warming (in accord with Bergmann's rule).
Location  Global.
Methods  We used > 4400 specimens representing 22 carnivore species in 52 populations collected over the last few decades to examine whether size changed with collection date when geography and sex are accounted for. We then examined several factors related to global warming, body mass, diet, and the attributes of the different datasets, to see whether they affect the standardized slope (β) of the size versus time regression.
Results  Six of 52 populations we examined show a significant effect of year of collection on body size at the 0.05 probability level. The response of size to global warming does not reflect spatial patterns of size variation, nor do diet or body mass affect tendency of populations to change in body size. Size changes are no more pronounced in populations that have been sampled more recently. However, change, where it occurs, is rapid.
Main conclusions  There may be a tendency in the literature to report only cases where recent changes are prevalent. Although in our data only a minority of populations show body size changes, we may see changes accelerating in the future in response to more drastic climatic changes and other anthropogenic changes.     

Mammalian island biogeography: effects of area,isolation and vagility

Summary In this paper I reviewed mammalian biogeography for 19 archipelagoes and tested the applicability of the equilibrium theory of island biogeography to mammalian faunas in general. The species-area and species-isolation relationships of terrestrial mammals were consistent with the basic predictions of the equilibrium theory. The z-values for the species-area relationship did not differ significantly from Preston's canonical value of 0.26 (P>0.50), and the modal z-value for non-volant mammals was 0.25. Moreover, z-values increased with isolation and decreased with vagility of the fauna in question. Furthermore, the strength of the species-isolation correlation was negatively correlated with island area and vagility (P<0.001).     

Global biogeography and ecology of body size in birds

In 1847, Karl Bergmann proposed that temperature gradients are the key to understanding geographic variation in the body sizes of warm-blooded animals. Yet both the geographic patterns of body-size variation and their underlying mechanisms remain controversial. Here, we conduct the first assemblage-level global examination of 'Bergmann's rule' within an entire animal class. We generate global maps of avian body size and demonstrate a general pattern of larger body sizes at high latitudes, conforming to Bergmann's rule. We also show, however, that median body size within assemblages is systematically large on islands and small in species-rich areas. Similarly, while spatial models show that temperature is the single strongest environmental correlate of body size, there are secondary correlations with resource availability and a strong pattern of decreasing body size with increasing species richness. Finally, our results suggest that geographic patterns of body size are caused both by adaptation within lineages, as invoked by Bergmann, and by taxonomic turnover among lineages. Taken together, these results indicate that while Bergmann's prediction based on physiological scaling is remarkably accurate, it is far from the full picture. Global patterns of body size in avian assemblages are driven by interactions between the physiological demands of the environment, resource availability, species richness and taxonomic turnover among lineages.     

Insular biogeography of mammals in Canadian parks: a re-analysis

Aim Glenn & Nudds (1989) compared mammal species richness in Canadian parks to estimated species-area relationships prior to European settlement to test if parks presently contain their historical compliment of species. However, the data they used to estimate the presettlement species-area relationships were not commensurate with the scale of the parks and were not independent. This uncertainty reduces the utility of Glenn and Nudds analysis to detect which, if any, parks are experiencing mammal extirpations and ultimately to direct conservation efforts to enhance mammal conservation in Canada. We improved Glenn and Nudds methods and re-assesed the conservation status of mammals in Canadian parks. Location Canada. Methods We constructed species-area curves for disturbance tolerant and intolerant species in five mammal regions of Canada by sampling historical range maps using plots of 10– 10,000 km². We compared these estimates of expected species richness to current species richness in thirty-six parks and contrast our results with those of Glenn and Nudds. Results All ten re-estimated species-area regressions had higher intercepts than those reported by Glenn and Nudds; four had lower slopes. Of seventy-two cases analysed, we found twenty in which parks had fewer species than expected and four in which parks had more species than expected, compared to fourteen and thirty-three, respectively, reported by Glenn and Nudds. Changes in both the regression parameters and increases in the prediction intervals accounted for the disparity between these results. Park residuals were not significantly different from zero in eight of ten analyses. Residuals for parks in the Alleghenian-Illinoian mammal province were negative and decreased with increasing park size. Main conclusions Improvement of Glenn and Nudds methods had an effect on both the parameter estimates and precison of the presettlement species-area curves. As a result, mammal conservation in some Canadian parks is worse than Glenn and Nudds reported. Six additional parks contained fewer disturbance intolerant species than predicted. However, Glenn and Nudds finding of systematic differences between expected and observed species richness in parks in the densely populated region of southern Ontario, southern Quebec and the maritime provinces was not an artefact of their methods. In this region, seven of ten parks appear to have lost disturbance-sensitive species of mammals. The pattern of species loss in this region was consistent with the idea that these parks have become isolated from mammal dispersal by surrounding habitat change and have experienced local species extirpations.     

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.     

Small island biogeography in the Gulf of California: lizards, the subsidized island biogeography hypothesis, and the small island effect

Aim We used insular lizard communities to test the predictions of two hypotheses that attempt to explain patterns of species richness on small islands. We first address the subsidized island biogeography (SIB) hypothesis, which predicts that spatial subsidies may cause insular species richness to deviate from species–area predictions, especially on small islands. Next, we examine the small island effect (SIE), which suggests small islands may not fit the traditional log‐linear species–area curve. Location Islands with arthropodivorous lizard communities throughout the Gulf of California. Methods To evaluate the SIB hypothesis, we first identified subsidized and unsubsidized islands based on surrogate measures of allochthonous productivity (i.e. island size and bird presence). Subsequently, we created species–area curves from previously published lizard species richness and island area data. We used the residuals and slopes from these analyses to compare species richness on subsidized and unsubsidized islands. To test for an SIE, we used breakpoint regression to model the relationship between lizard species richness and island area. We compared results from this model to results from the log‐linear regression model. Results Subsidized islands had a lower slope than unsubsidized islands, and the difference between these groups was significant when small islands were defined as < 1 km². In addition to comparing slopes, we tested for differences in the magnitude of the residuals (from the species–area regression of all islands) for subsidized vs. unsubsidized islands. We found no significant patterns in the residual values for small vs. large islands, or between islands with and without seabirds. The SIE was found to be a slightly better predictor of lizard species richness than the traditional log‐linear model. Main conclusions Predictions of the SIB hypothesis were partially supported by the data. The absence of a significant SIE may be a result of spatial subsidies as explained by the SIB hypothesis and data presented here. We conclude by suggesting potential scenarios to test for interactions between these two small island hypotheses. Future studies considering factors affecting species richness should examine the possible role of spatial subsidies, an SIE, or a synergistic effect of the two in data sets with small islands.     

Biogeography of body size in Pacific island birds

Many insular vertebrates have undergone rapid and dramatic changes in body size compared to their mainland counterparts. Here we explore the relationship between two well known patterns of island body size – the tendency for large‐bodied species to dwarf and small‐bodied species to get larger on islands, known as the “island rule”, and the scaling of maximum and minimum body size of island assemblages with island area. Drawing on both fossil and modern data, we examined the relationship between body size and island area in Pacific island birds, both within clades and at the island assemblage level. We found that the size of the smallest bird on each island decreased with island area while the maximum body size increased with island area. Similarly, within clades the body size of small‐bodied groups decreased and large‐bodied groups increased from small to large islands, consistent with the island rule. However, the magnitude of size change within clades was not sufficient to explain the overall scaling of maximum size with island area. Instead, the pattern was driven primarily by the evolution of very large, flightless birds on large islands. Human‐mediated extinctions on islands over the past few millennia severely impacted large, flightless birds, to the effect that this macroecological pattern has been virtually erased. After controlling for effects of biogeographic region and island area, we found island productivity to be the best predictor of maximum size in flightless birds. This result, and the striking similarities in maximum body size between flightless birds and island mammals, suggests a common energetic mechanism linking body size and landmass area in both the island rule and the scaling of island body size extremes.     

On being the right size: food-limited feedback on optimal body size

An insular population of white-tailed deer Odocoileus virginianus introduced in 1896 to predator-free Anticosti Island, Quebec, has caused long-term changes in the plant community. Food quality declined as did body weight. Although different parameters of reproduction changed, overall reproductive rates remained similar, thus maintaining deer density and promoting further change in habitat. These results show (i) long-term feedbacks on carrying capacity, (ii) the mechanism for reduction of body mass, and (iii) the lack of strong reduction in reproductive rates to regulate the population at high density, a feature of Eutherians. They are relevant to mechanisms determining the evolution of vertebrate body sizes.     

The frog filter: amphibian introduction bias driven by taxonomy,body size and biogeography

Aim Invasive species often exhibit a highly non‐random suite of traits relative to non‐invasive taxa, and these biases reflect strong selection at a series of steps along the invasion pathway. Here we investigate traits that are favoured in the first of these steps: the introduction of species outside their native geographic range. We use the global introduction of amphibians as our case study. Location Global. Methods We examined whether taxonomy, body size and attributes of the native geographic range predict the likelihood of an introduction event in amphibians. We then determined whether these characteristics differed between amphibian species that have been introduced intentionally and those that have been introduced unintentionally (i.e. stowaways). Results Comparisons of introduced and non‐introduced amphibians reveal significant biases with respect to taxonomic position (c. 50% of all introduced species originate from only 5% of all families), and characteristics of the native geographic range (most introduced species originate from the Northern Hemisphere, occupy a wide geographic range and are sympatric with high densities of humans). Many of these biases presumably reflect the ease and likelihood of capturing and transporting such animals. Additionally, intentionally introduced species are of larger than average body size, occupy higher elevations in their native ranges, and are more likely to originate from the Northern Hemisphere than unintentionally introduced species. Main conclusions Introduced amphibian species are not a random subset of the global amphibian fauna with respect to taxonomic affiliation and biogeography, and this restricts our ability to make robust generalizations regarding the ecological determinants of introduction success. Nevertheless, many of our findings are similar to those revealed by previous analyses of vertebrate and invertebrate introductions. Thus, our study suggests that biases in the anthropogenic mechanisms involved in transporting species around the globe are surprisingly consistent across broadly divergent taxa.     

A neglected facet of island biogeography: The role of internal spatial dynamics in area effects

The classical theory of island biogeography has as its basic variable the presence or absence of species on entire islands, and as its basic processes colonization and extinction rates on entire islands as functions of island area, distance, and so forth. Yet for many organisms with limited dispersal abilities, it may be more reasonable to consider larger islands as comprised of an ensemble of local populations coupled by within-island dispersal. Conceptual arguments and a simple patch occupancy model are used to examine the potential relevance of such internal spatial dynamics in explaining area effects, expressed via the probability that a species is present per unit area as a function of total island area. The model suggests that strong area effects depend on a rather fine balance between local colonization and extinction rates. A fruitful direction of future research should be the application of patch dynamic theory to classic island biogeographic questions and systems.     

Changes in otolith area: sensory area ratio with body size and depth

Synopsis The ratio between the sensory areas of the sulcus acusticus and the sagittal otolith (S : O ratio) in the species Merluccius capensis and M. paradoxus was analyzed using a digital image processing system. Sagittal growth in both species is negatively allometric with respect to total fish length but is more negative in the deeper-dwelling species, Merluccius paradoxus. In contrast, the sulcus acusticus undergoes a relative increase in size, that is, the S : O ratio increases with fish length. There was a clear relationship between the increase in the S : 0 ratio and depth.     

Genetic and phylogenetic consequences of island biogeography

Abstract.— Island biogeography theory predicts that the number of species on an island should increase with island size and decrease with island distance to the mainland. These predictions are generally well supported in comparative and experimental studies. These ecological, equilibrium predictions arise as a result of colonization and extinction processes. Because colonization and extinction are also important processes in evolution, we develop methods to test evolutionary predictions of island biogeography. We derive a population genetic model of island biogeography that incorporates island colonization, migration of individuals from the mainland, and extinction of island populations. The model provides a means of estimating the rates of migration and extinction from population genetic data. This model predicts that within an island population the distribution of genetic divergences with respect to the mainland source population should be bimodal, with much of the divergence dating to the colonization event. Across islands, this model predicts that populations on large islands should be on average more genetically divergent from mainland source populations than those on small islands. Likewise, populations on distant islands should be more divergent than those on close islands. Published observations of a larger proportion of endemic species on large and distant islands support these predictions.     

The effect of island area on body size in a primate species from the Sunda Shelf Islands

Aim  We examine the effect of island area on body dimensions in a single species of primate endemic to Southeast Asia, the long-tailed macaque ( Macaca fascicularis ). In addition, we test Allen's rule and a within-species or intraspecific equivalent of Bergmann's rule (i.e. Rensch's rule) to evaluate body size and shape evolution in this sample of insular macaques.
Location  The Sunda Shelf islands of Southeast Asia.
Methods  Body size measurements of insular macaques gathered from the literature were analysed relative to island area, latitude, maximum altitude, isolation from the mainland and other islands, and various climatic variables using linear regression.
Results  We found no statistically significant relationship between island area and body length or head length in our sample of insular long-tailed macaques. Tail length correlated negatively with island area. Head length and body length exhibited increases corresponding to increasing latitude, a finding seemingly consistent with the expression of Bergmann's rule within a single species. These variables, however, were not correlated with temperature, indicating that Bergmann's rule is not in effect. Tail length was not correlated with either temperature or increasing latitude, contrary to that predicted by Allen's rule.
Main conclusions  The island rule dictating that body size will covary with island area does not apply to this particular species of primate. Our study is consistent with results presented in the literature by demonstrating that skull and body length in insular long-tailed macaques do not, strictly speaking, conform to Rensch's rule. Unlike previous studies, however, our findings suggest that tail-length variation in insular macaques does not support Allen's rule.     

Insular shifts in body size of rice frogs in the Zhoushan Archipelago, China

1. Differences in body size between mainland and island populations have been reported for reptiles, birds and mammals. Despite widespread recognition of insular shifts in body size in these taxa, there have been no reports of such body size shifts in amphibians. 2. We provide the first evidence of an insular shift in body size for an amphibian species, the rice frog Rana limnocharis. We found significant increases in body size of rice frogs on most sampled islands in the Zhoushan archipelago when compared with neighbouring mainland China. 3. Large body size in rice frogs on islands was significantly related to increased population density, in both breeding and non-breeding seasons. Increases in rice frog density were significantly related to higher resource availability on islands. Increased resource availability on islands has led to higher carrying capacities, which has subsequently facilitated higher densities and individual growth rates, resulting in larger body size in rice frogs. We also suggest that large body size has evolved on islands, as larger individuals are competitively superior under conditions of harsh intraspecific competition at high densities. 4. Increases in body size in rice frogs were not related to several factors that have been implicated previously in insular shifts in body size in other taxa. We found no significant relationships between body size of rice frogs and prey size, number of larger or smaller frog species, island area or distance of islands from the mainland. 5. Our findings contribute to the formation of a broad, repeatable ecological generality for insular shifts in body size across a range of terrestrial vertebrate taxa, and provide support for recent theoretical work concerning the importance of resource availability for insular shifts in body size.     

Growth strategies and optimal body size in temperate pararginii butterflies

In temperate insects the evolution of growth strategies andthe optimal age and size at maturity will depend strongly onseasonal variation in temperature and other resources. However,compared to photoperiod, temperature itself is a relativelypoor predictor of seasonal change and timing decisions in insectsare often most strongly influenced by the photoperiod. HereI review the evolution of seasonal growth strategies in thebutterfly tribe Pararginii (Satyrinae: Nymphalidae) and relateit to life history theory. The results indicate that individuallarvae may adjust their growth trajectories in relation to informationon time horizons obtained from the photoperiod. The growth strategiescan be characterized by a set of state-dependent decision rulesthat specify how an individual should respond to its internalstate and external circumstances. These decision rules may alsoinfluence how individual growth change with a rise in temperature,showing that the standard expectation of increased growth rateswith increasing temperatures may not always be true. With lesstime available individual larvae increase growth rates and therebyachieve shorter development times, most often without any effectson final sizes. One reason for the apparent optimization ofgrowth rate seems to be that growing fast may incur costs thatlarvae developing under lower time limitations chose to avoid.The patterns of growth found in these and many other studiesare difficult to reconcile with common assumptions of what typicallydetermines optimal body size in insects. In particular it seemsas if there should be some costs of a large body size that,so far, have been poorly documented.