Body Size Clines Research Articles

Body size–climate relationships of European spiders

AbstractAim Geographic body size patterns of mammals and birds can be partly understood under the framework of Bergmann’s rule. Climatic influences on body size of invertebrates, however, appear highly variable and lack a comparable, generally applicable theoretical framework. We derived predictions for body size–climate relationships for spiders from the literature and tested them using three datasets of variable spatial extent and grain.Location Europe.Methods To distinguish climate from space, we compared clines in body size within three datasets with different degrees of co‐variation between latitude and climate. These datasets were: (1) regional spider faunas from 40 European countries and large islands; (2) local spider assemblages from standardized samples in 32 habitats across Europe; and (3) local spider assemblages from Central European habitats. In the latter dataset climatic conditions were determined more by habitat type than by geographic position, and therefore this dataset provided a non‐spatial gradient of various microclimates. Spider body size was studied in relation to latitude, temperature and water availability.Results In all three datasets the mean body size of spider assemblages increased from cool/moist to warm/dry environments. This increase could be accounted for by turnover from small‐bodied to large‐bodied spider families. Body size–climate relationships within families were inconsistent.Main conclusions Starvation resistance and accelerated maturation can be ruled out as explanations for the body size clines recorded, because they predict the inverse of the observed relationship between spider body size and temperature. The relationship between body size and climate was partly independent of geographic position. Thus, the restriction of large‐bodied spiders to their glacial refugia owing to dispersal limitations can be excluded. Our results are consistent with mechanisms invoking metabolic rate, desiccation resistance and community interactions to predict a decrease in body size from warm and dry to cool and moist conditions.

Population biology of the long-nosed potoroo (Potorous tridactylus) in the Southern Highlands of New South Wales

The population biology of the long-nosed potoroo (Potorous tridactylus) was investigated at Barren Grounds Nature Reserve and Budderoo National Park in the Southern Highlands of New South Wales (NSW). Both study areas are important conservation reserves for this threatened species, with a large gap north (~300 km) to the next known viable population on the mid-north coast of NSW at Mount Royal. Potoroos were live-trapped using bandicoot-sized cage traps, each set ~100 m apart along walking tracks and fire trails. Trapping was conducted each autumn and spring over five years to enumerate the local population of potoroos and to describe their morphometrics. The local long-nosed potoroos were larger in size than those recorded to the south on mainland Australia, but smaller than those in north-eastern NSW, supporting the concept of a latitudinal cline in body size. Sexual dimorphism was observed, with adult males having larger body weights, head lengths and pes lengths. Between one-third and two-thirds of all males and females were captured in only a single trapping session, indicative of low levels of survivorship and/or high levels of dispersal or transience. Males regularly overlapped at trap sites with females, more so than with other males, while females rarely overlapped at trap sites. Barren Grounds Nature Reserve supported a larger number of potoroos and a greater degree of home range overlap between individuals.

Size, shape, and the thermal niche of endotherms

A key challenge in ecology is to define species' niches on the basis of functional traits. Size and shape are important determinants of a species' niche but their causal role is often difficult to interpret. For endotherms, size and shape define the thermal niche through their interaction with core temperature, insulation, and environmental conditions, determining the thermoneutral zone (TNZ) where energy and water costs are minimized. Laboratory measures of metabolic rate used to describe TNZs cannot be generalized to infer the capacity for terrestrial animals to find their TNZ in complex natural environments. Here, we derive an analytical model of the thermal niche of an ellipsoid furred endotherm that accurately predicts field and laboratory data. We use the model to illustrate the relative importance of size and shape on the location of the TNZ under different environmental conditions. The interaction between body shape and posture strongly influences the location of the TNZ and the expected scaling of metabolic rate with size at constant temperature. We demonstrate that the latter relationship has a slope of approximately (1/2) rather than the commonly expected surface area/volume scaling of (2/3). We show how such functional traits models can be integrated with spatial environmental datasets to calculate null expectations for body size clines from a thermal perspective, aiding mechanistic interpretation of empirical clines such as Bergmann's Rule. The combination of spatially explicit data with biophysical models of heat exchange provides a powerful means for studying the thermal niches of endotherms across climatic gradients.

Density‐dependent polyphenism and geographic variation in size among two populations of lubber grasshoppers (Romalea microptera)

Abstract.1. Density‐dependent phase polyphenism occurs when changes in density during the juvenile stages result in a developmental shift from one phenotype to another. Density‐dependent phase polyphenism is common among locusts (Orthoptera: Acrididae).2. Previously, we demonstrated a longitudinal geographic cline in adult body size (western populations = small adults; eastern populations = large adults) in the eastern lubber grasshopper (Romalea microptera) in south Florida. As lubbers are confamilial with locusts, we hypothesised that the longitudinal size cline was partly due to density‐dependent phase polyphenism.3. We tested the effect of density, population, and density×population interaction on life‐history traits (pronotum length, mass, cumulative development time, growth rate) of, and proportion surviving to, each of the five instars and the adult stage in a 2 × 3 factorial laboratory experiment with two lubber populations, each reared from hatchling to adult at three different densities.4. The effect of density on life history and survival was independent of the effects of population on life history and survival. Higher densities led to larger adult sizes (pronotum, mass) and lower survivorship. The western population had smaller adult masses, fewer cumulative days to the adult stage, and higher survivorship than the eastern population.5. Our data suggest that lubber grasshoppers exhibit density‐dependent phase polyphenism initiated by the physical presence of conspecifics. However, the plastic response of adult size to density observed in the laboratory is not consistent with the relationship between phenotypes and adult density in the field. Genetic differences between populations observed in the laboratory could contribute to size and life‐history differences among lubber populations in the field.

Shifting latitudinal clines in avian body size correlate with global warming in Australian passerines

Intraspecific latitudinal clines in the body size of terrestrial vertebrates, where members of the same species are larger at higher latitudes, are widely interpreted as evidence for natural selection and adaptation to local climate. These clines are predicted to shift in response to climate change. We used museum specimens to measure changes in the body size of eight passerine bird species from south-eastern Australia over approximately the last 100 years. Four species showed significant decreases in body size (1.8-3.6% of wing length) and a shift in latitudinal cline over that period, and a meta-analysis demonstrated a consistent trend across all eight species. Southern high-latitude populations now display the body sizes typical of more northern populations pre-1950, equivalent to a 7 degrees shift in latitude. Using ptilochronology, we found no evidence that these morphological changes were a plastic response to changes in nutrition, a likely non-genetic mechanism for the pattern observed. Our results demonstrate a generalized response by eight avian species to some major environmental change over the last 100 years or so, probably global warming.

Ontogenetic Mechanisms Underlying a Geographic Size Cline in a Grasshopper, Romalea microptera

Abstract Geographic clines in body size have been described for many species, but relatively few investigations have tested hypotheses for the ontogenetic mechanisms maintaining geographic clines. We formalize and test the predictions for the role of ontogenetic mechanisms (e.g., hatching timing and size, juvenile developmental time, juvenile growth rate) in maintaining a longitudinal cline in adult body size of lubber grasshoppers [Romalea microptera (Beauvois)]. To obtain hatching timing and size, we collected eggs from wild females from several populations along the longitudinal gradient in 2 yr (2006 and 2007) and measured hatchling size and hatch date. To obtain juvenile developmental time and growth rates, we surveyed populations along the longitudinal gradient during 2 yr (2006 and 2007) and estimated developmental time and growth rates. We found the developmental time (hatching to fourth instar) and female growth rates increase from west to east along the cline. Patterns of hatching timing and hatching size were not consistent with the size cline. The size cline becomes evident in the third instar and is magnified in the fourth and fifth instars. Our data suggest that the size cline arises from some combination of clinal variation in developmental time and female growth rates; prolonged development and greater growth rates lead to larger mean size. Equally important, we found no evidence that differences in hatching time or size are ontogenetic causes of this cline. Our hypotheses for ontogenetic mechanisms producing an adult size cline should serve as a template for ecologists seeking to understand the ontogenetic basis of spatial variation in phenotypes.

Bergmann's idiosyncratic rule: a role for fecundity selection?

The negative relationship between temperature and geographical variation in body size, or Bergmann's rule, is among the most thoroughly studied ecogeographical rules, yet the pattern and process underlying it remain controversial. Bergmann's original observations were of body size clines among endotherms, but in the last 50 years there has been increasing recognition that both Bergmann's rule and its reverse occur in many ectotherm taxa. A new study of syngnathid fish by Wilson (2009) in this issue of Molecular Ecology sheds light on intriguing alternative mechanisms that may explain variation in the direction of body size clines across taxa. Wilson shows that Bergmann's rule is found in pipefish of the genus Syngnathus, but not in seahorses of the genus Hippocampus. His results suggest that polygamy in pipefish allows fecundity selection to favour large size at low temperatures, compensating for increases in brooding time.

Immune Function, Body Size, and Parasite Load in Lubber Grasshoppers

Immunity is an important biological property of organisms that protects them from parasites. Similarly, body size is one of the most important biological traits because almost all biological processes, from the cellular to the ecosystem level, scale with body size. Our goal was to determine the correlation between body size and immune function in different populations of the eastern lubber grasshopper (Romalea microptera) which differ in body size. Field data was collected on grasshopper location, size (thorax and femur lengths), and immune function (measured by melanization response). In accordance with previous work, we found a significant body size cline among populations of south Florida grasshoppers: on average, small adult grasshoppers are found in western populations whereas large grasshoppers are found in eastern populations. However, we did not find a significant relationship between body size and one measure of immune function, either within or across these populations. Future work should be directed at understanding when body size and immune function might or might not be correlated.

Microgeographic life history variation in an alpine caddisfly: plasticity in response to seasonal time constraints

Summary1. Temporally constrained environments, such as habitats with short growth seasons or short hydroperiods, cause potentially strong selection on life histories. Depending on the predictability of these events and the extent of spatial and temporal heterogeneity, local populations could become adapted either via a fixed phenotype or via life history plasticity in response to these environmental cues.2. We used a common garden experiment to investigate microgeographic variation in life history responses to combined changes in photoperiod (ambient/late) and hydroperiod (constant/drying) time constraint cues in an alpine caddisfly (Trichoptera). We compared six populations (three permanent/three temporary streams) originating from a small, alpine floodplain and which spanned an expected gradient in growth period duration (GPD) with distance from glaciers.3. We made two main predictions in relation to locally varying selection pressures: (i) populations nearest glaciers (shorter GPD and strongest time constraints) should have the fastest development rates and (ii) populations from permanent streams should be less able to respond to drying hydroperiods than populations from temporary streams.4. All populations and both sexes accelerated development in response to late season photoperiod cues. However, only permanent stream populations showed an increase in development time with increasing GPD, suggesting that other factors were influencing populations in temporary streams.5. Permanent stream populations showed countergradient variation (genetic and environmental influences were in opposition) in development time, and under‐compensation of growth rates resulted in a converse Bergmann cline in body size (smaller body size along gradients of declining season length). The extent of plasticity in response to hydroperiod, and the combined effects of both time constraints, differed between populations and sexes, but were not consistent among populations.6. Taken together, our results suggest adaptive plasticity in response to season length. The lack of a predictable pattern in response to hydroperiod may be due to gene flow or weak selection. We conclude that spatially structured populations can strongly differ in phenotypic plasticity even at microgeographic scales.

Latitudinal Body Size Clines in the ButterflyPolyommatus icarusare Shaped by Gene-Environment Interactions

The study of latitudinal body size clines can illuminate processes of local adaptation, but there is a need for an increased understanding of the relative roles of genetic variation, environmental effectstions or this reason, we combined an investigation of a museum collection of the common blue butterfly Polyommatus icarus (Rottemburg) (Lycaenidae: Polyommatini) from Sweden with a common-garden experiment in the laboratory, using strains reared from individuals collected from three different latitudes. Sizes of the field-collected butterflies tended to smoothly decrease northwards in a latitudinal cline, but suddenly increase at the latitude where the life cycle changes from two to one generations per year, hence allowing more time for this single generation. Further north, the size of the field-collected butterflies again decreased with latitude (with the exception of the northernmost collection sites). This is in accordance with the “converse Bergmann” pattern and with the “saw-tooth model” suggesting that insect size is shaped by season length and number of generations along latitudinal transects. In contrast, under laboratory conditions with a constant long day-length there was a different pattern, with the butterflies pupating at a higher mass when individuals originated from southern populations under time stress to achieve a second generation. This is indirect evidence for field patterns being shaped by end-of-season cues cutting development short, and also suggests counter-gradient variation, as butterflies from the time-stressed populations over-compensated for decreasing larval development time by increasing their growth rates, thus obtaining higher mass. Hence, we found support for both adaptive phenotypic plasticity and local genetic adaptation, with gene-environment interactions explaining the observed field patterns.

PHYLOGENETIC META-ANALYSIS

Meta-analysis is a powerful statistical technique that combines the results of independent studies to identify general trends. When the species under examination are not independent however, it is also necessary to incorporate phylogenetic information into the analysis. Unfortunately, current meta-analytic approaches cannot account for lack of independence resulting from shared evolutionary history, so a general solution to this problem is lacking. In this article, I derive a model for phylogenetic meta-analysis, so that data across studies may be summarized with evolutionary history explicitly incorporated. The approach takes advantage of common aspects of linear statistical models used by both meta-analysis and the phylogenetic comparative method, thereby allowing them to be analytically combined. In this manner, the correlation structure generated by phylogenetic history can be incorporated directly into the meta-analytic procedure. I illustrate the approach by examining the prevalence of body size clines in mammals. The approach is general, and can also be used to incorporate correlation structure among studies generated by other factors, such as spatial or temporal proximity, or environmental similarity. Therefore, this procedure provides a general statistical template for meta-analytic techniques that can account for attributes that generate nonindependence among studies. Implications of the phylogenetic meta-analysis are discussed.

AMPHIBIANS DO NOT FOLLOW BERGMANN'S RULE

The tendency for organisms to be larger in cooler climates (Bergmann's rule) is widely observed in endotherms, and has been reputed to apply to some ectotherms including amphibians. However, recent reports provide conflicting support for the pattern, questioning whether Bergmann's clines are generally present in amphibians. In this study, we measured 96,996 adult Plethodon from 3974 populations to test for the presence of Bergmann's clines in these salamanders. Only three Plethodon species exhibited a significant negative correlation between body size and temperature consistent with Bergmann's rule, whereas 37 of 40 species did not display a pattern consistent with this prediction. Further, a phylogenetic comparative analysis found no relationship between body size and temperature among species. A meta-analysis combining our data with the available data for other amphibian species revealed no support for Bergmann's rule at the genus (Plethodon), order (Caudata), or class (Amphibia) levels. Our findings strongly suggest that negative thermal body size clines are not common in amphibians, and we conclude that Bergmann's rule is not generally applicable to these taxa. Thus, evolutionary explanations of Bergmann's clines in other tetrapods need not account for unique life-history attributes of amphibians.

Geographic Variation in Body Size and Sexual Size Dimorphism of a Seed‐Feeding Beetle

Body size of many animals varies with latitude: body size is either larger at higher latitudes (Bergmann's rule) or smaller at higher latitudes (converse Bergmann's rule). However, the causes underlying these patterns are poorly understood. Also, studies rarely explore how sexual size dimorphism varies with latitude. Here we investigate geographic variation in body size and sexual size dimorphism of the seed-feeding beetle Stator limbatus, collected from 95 locations along a 38 degrees range in latitude. We examine 14 variables to test whether clines in environmental factors are adequate to explain geographic patterns of body size. We found that body size and sexual size dimorphism of S. limbatus varied considerably with latitude; beetles were smaller but more dimorphic at lower latitudes. Body size was not correlated with a gradient in mean temperature, contrary to the commonly accepted hypothesis that clines are produced by latitudinal gradients in temperature. Instead, we found that three factors were adequate to explain the cline in body size: clinal variation in host plant seed size, moisture (humidity), and seasonality (variance in humidity, precipitation, and temperature). We also found that the cline in sexual size dimorphism was partially explainable by a gradient in moisture, though moisture alone was not sufficient to explain the cline. Other ecological or environmental variables must necessarily contribute to differences in selection on male versus female body size. The main implications of our study are that the sexes differ in the magnitude of clinal variation in body size, creating latitudinal variation in sexual size dimorphism, and that clines in body size of seed beetles are likely influenced by variation in host seed size, water availability, and seasonality.

Geographic Variation in Body Size and Sexual Size Dimorphism of a Seed-Feeding Beetle

Body size of many animals varies with latitude: body size is either larger at higher latitudes (Bergmann’s rule) or smaller at higher latitudes (converse Bergmann’s rule). However, the causes underlying these patterns are poorly understood. Also, studies rarely explore how sexual size dimorphism varies with latitude. Here we investigate geographic variation in body size and sexual size dimorphism of the seed‐feeding beetle Stator limbatus, collected from 95 locations along a 38° range in latitude. We examine 14 variables to test whether clines in environmental factors are adequate to explain geographic patterns of body size. We found that body size and sexual size dimorphism of S. limbatus varied considerably with latitude; beetles were smaller but more dimorphic at lower latitudes. Body size was not correlated with a gradient in mean temperature, contrary to the commonly accepted hypothesis that clines are produced by latitudinal gradients in temperature. Instead, we found that three factors were adequate to explain the cline in body size: clinal variation in host plant seed size, moisture (humidity), and seasonality (variance in humidity, precipitation, and temperature). We also found that the cline in sexual size dimorphism was partially explainable by a gradient in moisture, though moisture alone was not sufficient to explain the cline. Other ecological or environmental variables must necessarily contribute to differences in selection on male versus female body size. The main implications of our study are that the sexes differ in the magnitude of clinal variation in body size, creating latitudinal variation in sexual size dimorphism, and that clines in body size of seed beetles are likely influenced by variation in host seed size, water availability, and seasonality.

Fitness variation in response to artificial selection for reduced cell area, cell number and wing area in natural populations of Drosophila melanogaster

BackgroundGenetically based body size differences are naturally occurring in populations of Drosophila melanogaster, with bigger flies in the cold. Despite the cosmopolitan nature of body size clines in more than one Drosophila species, the actual selective mechanisms controlling the genetic basis of body size variation are not fully understood. In particular, it is not clear what the selective value of cell size and cell area variation exactly is. In the present work we determined variation in viability, developmental time and larval competitive ability in response to crowding at two temperatures after artificial selection for reduced cell area, cell number and wing area in four different natural populations of D. melanogaster.ResultsNo correlated effect of selection on viability or developmental time was observed among all selected populations. An increase in competitive ability in one thermal environment (18°C) under high larval crowding was observed as a correlated response to artificial selection for cell size.ConclusionViability and developmental time are not affected by selection for the cellular component of body size, suggesting that these traits only depend on the contingent genetic makeup of a population. The higher larval competitive ability shown by populations selected for reduced cell area seems to confirm the hypothesis that cell area mediated changes have a relationship with fitness, and might be the preferential way to change body size under specific circumstances.

The island rule and the evolution of body size in the deep sea

AbstractAim Our goal is to test the generality of the island rule – a graded trend from gigantism in small‐bodied species to dwarfism in large‐bodied species – in the deep sea, a non‐insular but potentially analogous system.Location Shallow‐water and deep‐sea benthic habitats in the western Atlantic Ocean from the North to South Poles.Methods We conducted regression analyses of body size of deep‐sea gastropods species relative to their shallow‐water congeners using measurements from the Malacolog ver. 3.3.3 database.Results Our results indicate that, consistent with the island rule, gastropod genera with small‐bodied, shallow‐water species have significantly larger deep‐sea representatives, while the opposite is true for genera that are large‐bodied in shallow water. Bathymetric body size clines within the deep sea are also consistent with predictions based on the island rule.Main conclusions Like islands, the deep sea is characterized by low absolute food availability, leading us to hypothesize that the island rule is a result of selection on body size in a resource‐constrained environment. The body size of deep‐sea species tends to converge on an optimal size for their particular ecological strategy and habitat.

WHEN RENSCH MEETS BERGMANN: DOES SEXUAL SIZE DIMORPHISM CHANGE SYSTEMATICALLY WITH LATITUDE?

Bergmann's and Rensch's rules describe common large-scale patterns of body size variation, but their underlying causes remain elusive. Bergmann's rule states that organisms are larger at higher latitudes (or in colder climates). Rensch's rule states that male body size varies (or evolutionarily diverges) more than female body size among species, resulting in slopes greater than one when male size is regressed on female size. We use published studies of sex-specific latitudinal body size clines in vertebrates and invertebrates to investigate patterns equivalent to Rensch's rule among populations within species and to evaluate their possible relation to Bergmann's rule. Consistent with previous studies, we found a continuum of Bergmann (larger at higher latitudes: 58 species) and converse Bergmann body size clines (larger at lower latitudes: 40 species). Ignoring latitude, male size was more variable than female size in only 55 of 98 species, suggesting that intraspecific variation in sexual size dimorphism does not generally conform to Rensch's rule. In contrast, in a significant majority of species (66 of 98) male latitudinal body size clines were steeper than those of females. This pattern is consistent with a latitudinal version of Rensch's rule, and suggests that some factor that varies systematically with latitude is responsible for producing Rensch's rule among populations within species. Identifying the underlying mechanisms will require studies quantifying latitudinal variation in sex-specific natural and sexual selection on body size.

Body proportions in ancient Andeans from high and low altitudes

Living human populations from high altitudes in the Andes exhibit relatively short limbs compared with neighboring groups from lower elevations as adaptations to cold climates characteristic of high-altitude environments. This study compares relative limb lengths and proportions in pre-Contact human skeletons from different altitudes to test whether ecogeographic variation also existed in Andean prehistory. Maximum lengths of the humerus, radius, femur, and tibia, and femoral head breadth are measured in sex-specific groups of adult human skeletons (N = 346) from the central (n = 80) and the south-central (n = 123) Andean coasts, the Atacama Desert at 2,500 m (n = 102), and the southern Peruvian highlands at 2,000-3,800 m (n = 41). To test whether limb lengths vary with altitude, comparisons are made of intralimb proportions, limb lengths against body mass estimates derived from published equations, limb lengths against the geometric mean of all measurements, and principal component analysis. Intralimb proportions do not statistically differ between coastal groups and those from the Atacama Desert, whereas intralimb proportions are significantly shorter in the Peruvian highland sample. Overall body size and limb lengths relative to body size vary along an altitudinal gradient, with larger individuals from coastal environments and smaller individuals with relatively longer limbs for their size from higher elevations. Ecogeographic variation in relation to climate explains the variation in intralimb proportions, and dietary variation may explain the altitudinal cline in body size and limb lengths relative to body size. The potential effects of gene flow on variation in body proportions in Andean prehistory are also explored.

Effects of temperature on cell size and number in the yellow dung fly Scathophaga stercoraria

(1) A number of hypotheses suggest that the temperature-size rule (larger at cooler temperatures), and consequently Bergmann clines in whole-organism body size (larger at higher latitudes), may be a mere consequence of processes at the cellular level, i.e., a physiological constraint. (2) We show that in the yellow dung fly, Scathophaga stercoraria (Diptera: Scathophagidae), the temperature-size rule holds for wing cell and ommatidia size. Increases in cell number made up two-thirds (eye) to three-quarters (wing) of the increase in organ size. Temperature effects on body size can be fully explained by its effects on cell size and number. (3) Our study adds to the generality of previous results in Drosophila spp. The physiological constraint hypothesis remains viable as a proximate, non-adaptive explanation for the temperature-size rule in ectotherms.

Body Size Clines in Sceloporus Lizards: Proximate Mechanisms and Demographic Constraints

Although most species of animals examined to date exhibit Bergmann's clines in body size, squamates tend to exhibit opposing patterns. Squamates might exhibit reversed Bergmann's clines because they tend to behaviorally regulate their body temperature effectively; the outcome of this thermoregulation is that warmer environments enable longer daily and annual durations of activity than cooler environments. Lizards of the genus Sceloporus provide an opportunity to understand the factors that give rise to contrasting thermal clines in body size because S. undulatus exhibits a standard Bergmann's cline whereas S. graciosus exhibits a reverse Bergmann's cline. Interestingly, rapid growth by individuals of both species involves adjustments of physiological processes that enable more efficient use of food. Patterns of adult body size are likely the evolutionary consequence of variation in juvenile survivorship among populations. In S. undulatus, delayed maturation at a relatively large body size is exhibited in cooler environments where juveniles experience higher survivorship, resulting in a Bergmann's cline. In S. graciosus, high juvenile survivorship is not consistently found in cooler environments, resulting in no cline or a reversed Bergmann's cline, i.e., geographic patterns in body size aren't necessarily produced by natural selection. Thus, discerning the mechanistic links between the thermal physiology of an organism and environment-specific rates of mortality will be critical to understanding the evolution of body size in relation to environmental temperature.