Patterns of size variation in bees at a continental scale: does Bergmann's rule apply?

Body size latitudinal clines have been widley explained by the Bergmann's rule in homeothermic vertebrates. However, there is no general consensus in poikilotherms organisms in particular in insects that represent the large majority of wildlife. Among them, bees are a highly diverse pollinators group with high economic and ecological value. Nevertheless, no comprehensive studies of species assemblages at a phylogenetically larger scale have been carried out even if they could identify the traits and the ecological conditions that generate different patterns of latitudinal size variation. We aimed to test Bergmann's rule for wild bees by assessing relationships between body size and latitude at continental and community levels. We tested our hypotheses for bees showing different life history traits (i.e. sociality and nesting behaviour). We used 142 008 distribution records of 615 bee species at 50 × 50 km (CGRS) grids across the West Palearctic. We then applied generalized least squares fitted linear model (GLS) to assess the relationship between latitude and mean body size of bees, taking into account spatial autocorrelation. For all bee species grouped, mean body size increased with higher latitudes, and so followed Bergmann's rule. However, considering bee genera separately, four genera were consistent with Bergmann's rule, while three showed a converse trend, and three showed no significant cline. All life history traits used here (i.e. solitary, social and parasitic behaviour; ground and stem nesting behaviour) displayed a Bergmann's cline. In general there is a main trend for larger bees in colder habitats, which is likely to be related to their thermoregulatory abilities and partial endothermy, even if a ‘season length effect’ (i.e. shorter foraging season) is a potential driver of the converse Bergmann's cline particularly in bumblebees.     

Evolution of body size in food webs: does the energetic equivalence rule hold?

The energetic equivalence rule (EER), which is derived from empirical observations linking population density and body size and from the allometric law linking metabolism and body size, predicts that the amount of energy used by the various species should be independent of body size. Here, we examine this hypothesis using a model that allows entire food webs to emerge from coevolution of interacting species. Body size influences both individual metabolism and interactions among species in the model. Overall, population density does decrease with body size roughly following a power law whose exponent is variable. We discuss this variability in the light of empirical data sets. The emerging relationship between the flux of resources exploited by the various species and their body size follows a decreasing power law, thus contradicting the EER. Our model emphasizes the importance of considering the influence of body size on species interactions in attempting to explain large-scale patterns related to body size.     

Cope's rule in cryptodiran turtles: do the body sizes of extant species reflect a trend of phyletic size increase?

Cope's rule of phyletic size increase is questioned as a general pattern of body size evolution. Most studies of Cope's rule have examined trends in the paleontological record. However, neontological approaches are now possible due to the development of model-based comparative methods, as well as the availability of an abundance of phylogenetic data. I examined whether the phylogenetic distribution of body sizes in extant cryptodiran turtles is consistent with Cope's rule. To do this, I examined body size evolution in each of six major clades of cryptodiran turtles and also across the whole tree of cryptodirans (n = 201 taxa). Extant cryptodiran turtles do not appear to follow Cope's rule, as no clade showed a significant phyletic body size trend. Previous analyses in other extant vertebrates have also found no evidence for phyletic size increase, which is in contrast to the paleontological data that support the rule in a number of extinct vertebrate taxa.     

Phenotypic plasticity of abdomen pigmentation in two geographic populations of Drosophila melanogaster: male–female comparison and sexual dimorphism

In Drosophila melanogaster male, the last abdominal tergites (A5–A6) are completely dark due to a strong internal constraint while, in female, all abdominal tergites (A2–A7) are phenotypically variable and highly plastic. Male A2–A4 are quite similar to those of female, but their plasticity was never investigated. In this paper, we compared the phenotypic plasticity of A2–A4 in both sexes in order to know if the major dimorphism (SD) expressed in male A5–A6 also extended toward the more anterior segments. We also compared two geographic populations living under very different climates in order to know if adaptive differences, previously observed in females also existed in males. With an isofemale line design, pigmentation variation according to growth temperature was investigated in the two populations from France and India. Male and female data were compared and sexual dimorphism (SD) analyzed in various ways. Reaction norms were quite similar in both sexes for A2 and A3, but clearly different for A4. Considering the total pigmentation (A2 + A3 + A4) males were darker than females at low temperatures and either identical to them (France) or lighter (India) above 25°C. SD (male–female difference) was genetically variable among lines and significantly different among segments. Reaction norms of SD exhibited an overall decrease with temperature and also a significant difference among populations, suggesting a local adaptation of SD to thermal conditions. The three plastic segments in male (A2–A4) seem to react adaptively to the thermal environment more efficiently than the same segments in female, in agreement with the thermal budget hypothesis. To our knowledge, it is the first time that a SD trait exhibits an adaptive difference between geographic populations.     

The temperature–size rule in a rotifer is determined by the mother and at the egg stage

In most ectotherms, compared with development at low temperatures, development at high temperatures results in the acceleration of maturation, which in turn results in a smaller size (temperature–size rule, TSR). It is not known at which developmental stages this thermal response is determined. We exposed different life stages of the rotifer Lecane inermis to 15, 20, or 25 °C to determine whether the TSR in the F1 generation is governed by the thermal conditions experienced by the mothers (F0 generation) during their development, during egg production, or during the development of the eggs or hatchlings. We found that the adult size was affected by the thermal conditions experienced by the mothers and embryos, but not by the conditions during post-hatching growth. We suggest that the thermal plasticity producing the TSR in rotifers may reflect the joint impacts of a maternal effect and a direct effect of the environment during egg development. The two-point control of the TSR resembles the thermal determination of other biological phenomena, similar to the thermally determined sex determination in ectotherms. Our results contribute not only to better understanding the proximate mechanisms of TSR, but also to comprehending the general biological mechanisms of response to temperature, which is one of the most important ecological factors.     

Body size and elevation: do Bergmann's and Rensch's rule apply in the polytypic bushcricket Poecilimon veluchianus?

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Scaling of species diversity and body mass in mammals: Cope’s rule and the evolutionary cost of large size

Abstract

Equations are constructed describing the inverse correlation of species diversity and body mass in extant and Cenozoic mammals. Cope’s rule, the tendency for many mammal clades to increase in body size through time, through phyletic change in single lineages or turnover within species groups, is interpreted as a probability function reducing diversity potential as a tradeoff for ecological/evolutionary gains. The inverse rule predicts that large species in clades will be less diverse than smaller species and, unless origination rates remain high among smaller clade members, clades conforming to Cope’s rule will decline in diversity, moving towards extinction. This proposition is evaluated in the Cenozoic histories of five North American mammal clades; cotton rats, felids, canids, hyaenodontids, and equids. Diversity potential of different size classes within the 3.75 million year phyletic history of the muskrat, Ondatra zibethicus, is also examined. A corollary prediction of the inverse rule, that large species should have longer durations (species lifespans) than small species, is unresolved. Successful clades maintain small size or a significant number of smaller species relative to clade average size. The potential loss of unique extant large mammal species justifies the conservation effort to protect them. The similarity of scaling exponents of species diversity to mass around a slope of -1.0 suggests that species diversity is correlated with home range size, the latter related to the probability of population fragmentation.     

Genetic estimates of contemporary effective population size: to what time periods do the estimates apply?

Although most genetic estimates of contemporary effective population size (Ne) are based on models that assume Ne is constant, in real populations Ne changes (often dramatically) over time, and estimates (Ne) will be influenced by Ne in specific generations. In such cases, it is important to properly match Ne to the appropriate time periods (for example, in computing Ne/N ratios). Here I consider this problem for semelparous species with two life histories (discrete generations and variable age at maturity--the 'salmon' model), for two different sampling plans, and for estimators based on single samples (linkage disequilibrium, heterozygote excess) and two samples (temporal method). Results include the following. Discrete generations: (i) Temporal samples from generations 0 and t estimate the harmonic mean Ne in generations 0 through t - 1 but do not provide information about Ne in generation t; (ii) Single samples provide an estimate of Ne in the parental generation, not the generation sampled; (iii) single-sample and temporal estimates never provide information about Ne in exactly the same generations; (iv) Recent bottlenecks can downwardly bias estimates based on linkage disequilibrium for several generations. Salmon model: (i) A pair of single-cohort (typically juvenile) samples from years 0 and t provide a temporal estimate of the harmonic mean of the effective numbers of breeders in the two parental years (N b(0) and N b(t)), but adult samples are more difficult to interpret because they are influenced by Nb in a number of previous years; (ii) For single-cohort samples, both one-sample and temporal methods provide estimates of Nb in the same years (contrast with results for discrete generation model); (iii) Residual linkage disequilibrium associated with past population size will not affect single-sample estimates of Nb as much as in the discrete generation model because the disequilibrium diffuses among different years of breeders. These results lead to some general conclusions about genetic estimates of Ne in iteroparous species with overlapping generations and identify areas in need of further research.     

Phenotypic plasticity in life‐history traits of Daphnia galeata in response to temperature – a comparison across clonal lineages separated in time

Climatic changes are projected to result in rapid adaptive events with considerable phenotypic shifts. In order to reconstruct the impact of increased mean water temperatures during past decades and to reveal possible thermal micro‐evolution, we applied a resurrection ecology approach using dormant eggs of the freshwater keystone species Daphnia galeata. To this end, we compared the adaptive response of D. galeata clones from Lake Constance of two different time periods, 1965–1974 (“historical”) versus 2000–2009 (“recent”), to experimentally increased temperature regimes. In order to distinguish between genetic versus environmentally induced effects, we performed a common garden experiment in a flow‐through system and measured variation in life‐history traits. Experimental thermal regimes were chosen according to natural temperature conditions during the reproductive period of D. galeata in Central European lakes, with one additional temperature regime exceeding the currently observable maximum (+2°C). Increased water temperatures were shown to significantly affect measured life‐history traits, and significant “temperature × clonal age” interactions were revealed. Compared to historical clones, recent clonal lineages exhibited a shorter time to first reproduction and a higher survival rate, which may suggest temperature‐driven micro‐evolution over time but does not allow an explicit conclusion on the adaptive nature of such responses.     

From cells to colonies: at what levels of body organization does the ‘temperature‐size rule’ apply?

SUMMARY An inverse relationship between temperature during ontogeny and final body size is widespread in ectotherms, but poorly understood. Evidence suggests that within organs, this “temperature‐size rule” (TSR) may also apply to cell size with no change in numbers. So how closely do reductions in size and number of cells and other repeated structures correlate with size reduction at higher levels of organization? We examine this in the context of a proposal that size and/or number changes at various organizational levels are adaptive responses to temperature‐ and size‐dependent oxygen supply. We subjected two clones of the modular colonial bryozoan, Celleporella hyalina, to orthogonal combinations of two temperatures and two oxygen concentrations during ontogeny, observing effects on sizes of colonies and larvae, and sizes and numbers of cells, tentacles, and modules (autozooids). We found that the size:number responses varied among cell types and among structures at different levels of organization, with the inverse temperature‐size relationship applying only to larval parenchymal cells and colony modules. Using our findings and other evidence we propose a unifying adaptive hypothesis that predicts how temperature affects the sizes of mitochondria, cells, organs, modules and organisms, and their relationships with processes that determine the functional capacity of aerobic metabolism.     

Cell and leaf size plasticity in Arabidopsis: what is the role of endoreduplication?

Leaf area expansion is affected by environmental conditions because of differences in cell number and/or cell size. Increases in the DNA content (ploidy) of a cell by endoreduplication are related to its size. The aim of this work was to determine how cell ploidy interacts with the regulation of cell size and with leaf area expansion. The approach used was to grow Arabidopsis thaliana plants performing increased or decreased rounds of endoreduplication under shading and water deficit. The shading and water deficit treatments reduced final leaf area and cell number; however, cell area was increased and decreased, respectively. These differences in cell size were unrelated to alterations of the endocycle, which was reduced by these treatments. The genetic modification of the extent of endoreduplication altered leaf growth responses to shading and water deficit. An increase in the extent of endoreduplication in a leaf rendered it more sensitive to the shade treatment but less sensitive to water deficit conditions. The link between the control of whole organ and individual cell expansion under different environmental conditions was demonstrated by the correlation between the plasticity of cell size and the changes in the duration of leaf expansion.     

Some mathematical aspects of chemotherapy—II: The distribution of a drug in the body

In a previous paper (Bellman, Jacquez, and Kalaba,Bull. Math. Biophysics,22: 181–198, 1960) a model of the processes occurring in the exchange of a drug between capillary plasma, extracellular space and intracellular space was developed. This included the possibility of a reaction between the drug and a component of the intracellular space. The equations developed thus describe the events within a capillary bed. In the present paper, a simplified model of the body is set up. Each organ is treated as a single capillary bed and is linked to other organs via the circulation, in the parallel and/or series arrangements found in the body. Mixing in the circulation is included at the simplest possible level. The concentration of drug entering any one capillary bed is determined by the concentrations leaving all other capillary beds, the time lags, and mixing involved in the circulation. The equations describing these processes in conjunction with the equations of the processes occurring within each capillary bed lead to a large set of differential-difference equations.     

Is bigger really bigger? Differential responses to temperature in measures of body size of the mosquito,Aedes albopictus

When confronted with variation in temperature, most ectotherms conform to a growth rule that "hotter is smaller". This phenomenon can have important implications on population dynamics, interactions with other species, and adaptation to new environments for arthropods. However, the impact of other environmental factors and genetics may affect that general rule. Furthermore, most studies measure a single body part, and do not examine how temperature and other factors alter the allometric relationship between measurements of growth. In this study, we test the hypothesis that temperature and nutrition, while strongly affecting growth, do not change the allometric relationship between body mass and wing length in the mosquito Aedes albopictus. We tested this hypothesis by growing larval mosquitoes from two populations at five temperatures with three food levels. Contrary to our hypothesis, we find that temperature has a profound effect on allometry, with higher temperatures resulting in mosquitoes with shorter wings and greater body mass, and that the effects of temperature are dependent upon available food and population origin. We therefore reject our hypothesis and propose several testable mechanisms that will provide greater insight into the relationship between temperature, food, and measures of growth.     

Proximate causes of Rensch's rule: does sexual size dimorphism in arthropods result from sex differences in development time?

A prominent interspecific pattern of sexual size dimorphism (SSD) is Rensch's rule, according to which male body size is more variable or evolutionarily divergent than female body size. Assuming equal growth rates of males and females, SSD would be entirely mediated, and Rensch's rule proximately caused, by sexual differences in development times, or sexual bimaturism (SBM), with the larger sex developing for a proportionately longer time. Only a subset of the seven arthropod groups investigated in this study exhibits Rensch's rule. Furthermore, we found only a weak positive relationship between SSD and SBM overall, suggesting that growth rate differences between the sexes are more important than development time differences in proximately mediating SSD in a wide but by no means comprehensive range of arthropod taxa. Except when protandry is of selective advantage (as in many butterflies, Hymenoptera, and spiders), male development time was equal to (in water striders and beetles) or even longer than (in drosophilid and sepsid flies) that of females. Because all taxa show female-biased SSD, this implies faster growth of females in general, a pattern markedly different from that of primates and birds (analyzed here for comparison). We discuss three potential explanations for this pattern based on life-history trade-offs and sexual selection.     

Genetics of recent habitat contraction and reduction in population size: does isolation by distance matter?

Fragmentation and loss of natural habitats are recognized as major threats to contemporary flora and fauna. Detecting past or current reductions in population size is therefore a major aim in conservation genetics. Statistical methods developed to this purpose have tended to ignore the effects of spatial population structure. However in many species, individual dispersal is restricted in space and fine-scale spatial structure such as isolation by distance (IBD) is commonly observed in continuous populations. Using a simulation-based approach, we investigated how comparative and single-point methods, traditionally used in a Wright-Fisher (WF) population context for detecting population size reduction, behave for IBD populations. We found that a complex 'quartet' of factors was acting that includes restricted dispersal, population size (i.e. habitat size), demographic history, and sampling scale. After habitat reduction, IBD populations were characterized by a stronger inertia in the loss of genetic diversity than WF populations. This inertia increases with the strength of IBD, and decreases when the sampling scale increases. Depending on the method used to detect a population size reduction, a local sampling can be more informative than a sample scaled to habitat size or vice versa. However, IBD structure led in numerous cases to incorrect inferences on population demographic history. The reanalysis of a real microsatellite data set of skink populations from fragmented and intact rainforest habitats confirmed most of our simulation results.     

Testing the generality of the ‘leafing intensity premium’ hypothesis in temperate broad-leaved forests: a survey of variation in leaf size within and between habitats

Because leaf size scales negatively and isometrically with leaf number per shoot size (leafing intensity) in woody species, and because most tree and shrub species have small leaves, Kleiman and Aarssen (J Ecol 95:376–382, 2007) recently proposed that natural selection favors high leafing intensity resulting in small leaves, i.e., the leafing-intensity-premium hypothesis. However, empirical evidence for or against this hypothesis is still lacking. In addition, this hypothesis has not been examined in the context of how leaf size varies among habitats. To fill this void, we investigated leaf size frequency distributions of woody species from temperate China and explored the relationships among leaf mass, leaf number, and stem mass of current-year shoots of 133 woody species at low and high altitudes of three mountain ranges. The scaling relationships between leaf size and leafing intensity (leaf number per stem mass) were determined using both standardized major axis regression analyses and phylogenetically independent comparative techniques. In light of the leafing-intensity-premium hypothesis, we made three predictions: (1) leaf size frequency distributions should be right-skewed for each local study area and for the entire study region, (2) leafing intensities at different altitudes at different sites should differ while leafing intensities at comparable altitudes should be similar baring large taxonomic differences among sites, and (3) that leafing intensity should be higher for any given leaf size in habitats with small-leaved species. Significant negative and isometric scaling relationships between leaf size and leafing intensity were found to be consistently conserved independent of habitat type, both across species and across correlated evolutionary divergences. Within each mountain range or across the entire study region, leaf size frequency distributions were right-skewed, in accordance with our prediction. However, leafing intensity was smaller for any given leaf size at the altitude with smaller leafed species than for altitudes characterized by large leafed species, i.e., altitudes characterized by species with small leaves did not have consistently higher leafing intensities than other altitudes on each mountain range. Our analyses therefore indicate the direct adaptive value of leaf size but not the selective advantage in high leafing intensity as posited by the leafing-intensity-premium hypothesis. We suggest that this hypothesis explains less about the variation of leaf size among different habitats as it does about variation within habitats, i.e., the relative importance of the adaptive significance of leafing intensity and leaf size can and does vary with habitats.     

Why does the unimodal species richness–productivity relationship not apply to woody species: a lack of clonality or a legacy of tropical evolutionary history?

Aim  To study how differences in species richness patterns of woody and herbaceous plants may be influenced by ecological and evolutionary factors. Unimodal species richness–productivity relationships (SRPRs) have been of interest to ecologists since they were first described three decades ago for British herbaceous vegetation by J. P. Grime. The decrease in richness at high productivity may be due to competitive exclusion of subordinate species, or diverse factors related to evolution and dispersal. Unimodal SRPRs are most often reported for plants, but there are exceptions. For example, unimodal SRPRs are common in the temperate zone but not in the tropics. Similarly, woody species and forest communities in the Northern Hemisphere do not tend to show unimodal SRPRs.
Location  Global.
Methods  We used data from the literature to test whether a unimodal SRPR applies to woody species and forest communities on a global scale. We explored whether the shape of SRPRs may be related to the lack of clonality in woody species (which may prevent their being competitively superior), or the legacy of evolutionary history (most temperate woody species originate from tropical lineages, and due to niche conservatism they may still demonstrate 'tropical patterns'). We used case studies that reported the names of the dominant or most abundant species for productive sites.
Results  Woody species were indeed less clonal than herbaceous species. Both clonality and the temperate evolutionary background of dominating species were associated with unimodality in SRPRs, with woodiness modifying the clonality effect.
Main conclusions  The unimodal SRPR has been common in the ecological literature because most such studies originate from temperate herbaceous communities with many clonal species. Consequently, both evolutionary and ecological factors may influence species richness patterns.     

Variation in the daily rhythm of body temperature of free-living Arabian oryx (Oryx leucoryx): does water limitation drive heterothermy?

Heterothermy, a variability in body temperature beyond the limits of homeothermy, has been advanced as a key adaptation of Arabian oryx (Oryx leucoryx) to their arid-zone life. We measured body temperature using implanted data loggers, for a 1-year period, in five oryx free-living in the deserts of Saudi Arabia. As predicted for adaptive heterothermy, during hot months compared to cooler months, not only were maximum daily body temperatures higher (41.1 ± 0.3 vs. 39.7 ± 0.1°C, P = 0.0002) but minimum daily body temperatures also were lower (36.1 ± 0.3 vs. 36.8 ± 0.2°C, P = 0.04), resulting in a larger daily amplitude of the body temperature rhythm (5.0 ± 0.5 vs. 2.9 ± 0.2°C, P = 0.0007), while mean daily body temperature rose by only 0.4°C. The maximum daily amplitude of the body temperature rhythm reached 7.7°C for two of our oryx during the hot-dry period, the largest amplitude ever recorded for a large mammal. Body temperature variability was influenced not only by ambient temperature but also water availability, with oryx displaying larger daily amplitudes of the body temperature rhythm during warm-dry months compared to warm-wet months (3.6 ± 0.6 vs. 2.3 ± 0.3°C, P = 0.005), even though ambient temperatures were the same. Free-living Arabian oryx therefore employ heterothermy greater than that recorded in any other large mammal, but water limitation, rather than high ambient temperature, seems to be the primary driver of this heterothermy.