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Recent studies have shown that most tetrapod groups (mammals, birds, chelonians, amphibians) show general intraspecific tendencies for increasing body size with latitude, whereas squamates (lizards and snakes) show an intraspecific tendency towards decreasing body size with latitude. Here I evaluate whether these size trends are general by using independent contrasts analysis to investigate the dependence of intraspecific size-latitude relationships (r), and the magnitude alone of size-latitude relationships ([r]), for tetrapod vertebrates, on sample size, range of latitudes sampled, average latitude sampled, and body size. Range of latitudes sampled, average latitude sampled, and body size did not influence body size-latitude relationships (r) or the magnitude alone of body size-latitude relationship ([r]). Sample size did not influence size-latitude relationships (r), but did influence the magnitude alone of size-latitude relationships ([r]), possibly indicating increased precision of estimating size-latitude relationships with increased sampling. In short, intraspecific size-latitude relationships are similar for species of different sizes, occurring at different latitudes, sampled over different latitudinal ranges, and differing in number of populations sampled (though magnitude alone is influenced by sample size). These results suggest that intraspecific size-latitude trends are general, and biologically significant (i.e., are not artifacts of sampling), thus deserving explanation.
Two seemingly opposite evolutionary patterns of clinal variation in body size and associated life history traits exist in nature. According to Bergmann's rule, body size increases with latitude, a temperature effect. According to the converse Bergmann rule, body size decreases with latitude, a season length effect. A third pattern causally related to the latter is countergradient variation, whereby populations of a given species compensate seasonal limitations at higher latitudes by evolving faster growth and larger body sizes compared to their low latitude conspecifics. We discuss these patterns and argue that they are not mutually exclusive because they are driven by different environmental causes and proximate mechanisms; they therefore can act in conjunction, resulting in any intermediate pattern. Alternatively, Bergmann and converse Bergmann clines can be interpreted as over- and undercompensating countergradient variation, respectively. We illustrate this with data for the wide-spread yellow dung fly, Scathophaga stercoraria (Diptera: Scathophagidae), which in Europe shows a Bergmann cline for size and a converse Bergmann cline (i.e., countergradient variation) for development time. A literature review of the available evidence on arthropod latitudinal clines further shows a patterned continuum of responses. Converse Bergmann clines due to end-of-season time limitations are more common in larger species with longer development times. Our study thus provides a synthesis to the controversy about the importance of Bergmann's rule and the converse Bergmann rule in nature.
Bivalve molluscs have a highly plastic feeding and growth physiology. The increasing availability of families artificially selected for faster growth has enabled physiological experiments to investigate the genetic basis for variable rates of growth. Fast growth is achieved by a combination of increased rates of feeding, reduced metabolic rates and lower metabolic costs of growth. In at least one species there is a trade-off between growth in protein and the storage of lipids that are utilized in gametogenesis. Energy requirements for maintenance are also higher in slow-growing individuals. Reduced costs of growth are due in part to increased efficiencies of protein turnover. Nevertheless, high protein turnover (and therefore high metabolic cost) may benefit fitness in the later stages of gametogenesis. Faster feeding rates do not impair flexibility in feeding behavior which compensates for changes in the food environment. Both inter- and intra-species differences in feeding behavior are evident and suggest possible constraints imposed by faster feeding on the efficiency of selection between food particles of different nutritional value.
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.
The general effects of temperature and nutritional quality on growth rate and body size are well known. We know little, however, about the physiological mechanisms by which an organism translates variation in diet and temperature into reaction norms of body size or development time. We outline an endocrine-based physiological mechanism that helps explain how this translation occurs in the holometabolous insect Manduca sexta (Sphingidae). Body size and development time are controlled by three factors: (i) growth rate, (ii) the timing of the cessation of juvenile hormone secretion (measured by the critical weight) and (iii) the timing of ecdysteroid secretion leading to pupation (the interval to cessation of growth [ICG] after reaching the critical weight). Thermal reaction norms of body size and development time are a function of how these three factors interact with temperature. Body size is smaller at higher temperatures, because the higher growth rate decreases the ICG, thereby reducing the amount of mass that can accumulate. Development time is shorter at higher temperatures because the higher growth rate decreases the time required to attain the critical weight and, independently, controls the duration of the ICG. Life history evolution along altitudinal, latitudinal and seasonal gradients may occur through differential selection on growth rate and the duration of the two independently controlled determinants of the growth period.
Ectothermic animals exhibit two distinct kinds of plasticity in response to temperature: Thermal performance curves (TPCs), in which an individual's performance (e.g., growth rate) varies in response to current temperature; and developmental reaction norms (DRNs), in which the trait value (e.g., adult body size or development time) of a genotype varies in response to developmental temperatures experienced over some time period during development. Here we explore patterns of genetic variation and selection on TPCs and DRNs for insects in fluctuating thermal environments. First, we describe two statistical methods for partitioning total genetic variation into variation for overall size or performance and variation in plasticity, and apply these methods to available datasets on DRNs and TPCs for insect growth and size. Our results indicate that for the datasets we considered, genetic variation in plasticity represents a larger proportion of the total genetic variation in TPCs compared to DRNs, for the available datasets. Simulations suggest that estimates of the genetic variation in plasticity are strongly affected by the number and range of temperatures considered, and by the degree of nonlinearity in the TPC or DRN. Second, we review a recent analysis of field selection studies which indicates that directional selection favoring increased overall size is common in many systems—that bigger is frequently fitter. Third, we use a recent theoretical model to examine how selection on thermal performance curves relates to environmental temperatures during selection. The model predicts that if selection acts primarily on adult size or development time, then selection on thermal performance curves for larval growth or development rates is directly related to the frequency distribution of temperatures experienced during larval development. Using data on caterpillar temperatures in the field, we show that the strength of directional selection on growth rate is predicted to be greater at the modal (most frequent) temperatures, not at the mean temperature or at temperatures at which growth rate is maximized. Our results illustrate some of the differences in genetic architecture and patterns of selection between thermal performance curves and developmental reaction norms.
Drosophila subobscura is a European (EU) species that was introduced into South America (SA) approximately 25 years ago. Previous studies have found rapid clinal evolution in wing size and in chromosome inversion frequency in the SA colonists, and these clines parallel those found among the ancestral EU populations. Here we examine thermoplastic changes in wing length in flies reared at 15, 20, and 25°C from 10 populations on each continent. Wings are plastically largest in flies reared at 15°C (the coldest temperature) and genetically largest from populations that experience cooler temperatures on both continents. We hypothesize that flies living in cold temperatures benefit from reduced wing loading: ectotherms with cold muscles generate less power per wing beat, and hence larger wings and/or a smaller mass would facilitate fight. We develop a simple null model, based on isometric growth, to test our hypothesis. We find that both EU and SA flies exhibit adaptive plasticity in wing loading: flies reared at 15°C generally have lower wing loadings than do flies reared at 20°C or 25°C. Clinal patterns, however, are strikingly different. The ancestral EU populations show adaptive clinal variation at rearing a temperature of 15°C: flies from cool climates have lower wing loadings. In the colonizing populations from SA, however, we cannot reject the null model: wing loading increases with decreasing clinal temperatures. Our data suggest that selective factors other than flight have favored the rapid evolution of large overall size at low environmental temperatures. However, selection for increased flight ability in such environments may secondarily favor reduced body mass.
In temperate insects the evolution of growth strategies and the optimal age and size at maturity will depend strongly on seasonal variation in temperature and other resources. However, compared to photoperiod, temperature itself is a relatively poor predictor of seasonal change and timing decisions in insects are often most strongly influenced by the photoperiod. Here I review the evolution of seasonal growth strategies in the butterfly tribe Pararginii (Satyrinae: Nymphalidae) and relate it to life history theory. The results indicate that individual larvae may adjust their growth trajectories in relation to information on time horizons obtained from the photoperiod. The growth strategies can be characterized by a set of state-dependent decision rules that specify how an individual should respond to its internal state and external circumstances. These decision rules may also influence how individual growth change with a rise in temperature, showing that the standard expectation of increased growth rates with increasing temperatures may not always be true. With less time available individual larvae increase growth rates and thereby achieve shorter development times, most often without any effects on final sizes. One reason for the apparent optimization of growth rate seems to be that growing fast may incur costs that larvae developing under lower time limitations chose to avoid. The patterns of growth found in these and many other studies are difficult to reconcile with common assumptions of what typically determines optimal body size in insects. In particular it seems as if there should be some costs of a large body size that, so far, have been poorly documented.
Basically all organisms can be classified as determinate growers if their growth stops or almost stops at maturation, or indeterminate growers if growth is still intense after maturation. Adult size for determinate growers is relatively well defined, whereas in indeterminate growers usually two measures are used: size at maturation and asymptotic size. The latter term is in fact not a direct measure but a parameter of a specific growth equation, most often Bertalanffy's growth curve. At a given food level, the growth rate in determinate growers depends under given food level on physiological constraints as well as on investments in repair and other mechanisms that improve future survival. The growth rate in indeterminate growers consists of two phases: juvenile and adult. The mechanisms determining the juvenile growth rate are similar to those in determinate growers, whereas allocation to reproduction (dependent on external mortality rate) seems to be the main factor limiting adult growth. Optimal resource allocation models can explain the temperature-size rule (stating that usually ectotherms grow slower in cold but attain larger size) if the exponents of functions describing the size-dependence of the resource acquisition and metabolic rates change with temperature or mortality increases with temperature. Emerging data support both assumptions. The results obtained with the aid of optimization models represent just a rule and not a law: it is possible to find the ranges of production parameters and mortality rates for which the temperature-size rule does not hold.
Body size and temperature are the two most important variables affecting nearly all biological rates and times, especially individual growth or production rates. By favoring an optimal maturation age and reproductive allocation, natural selection links individual growth to the mortality schedule. A recent model for evolution of life histories for species with indeterminate growth, which includes most fish, successfully predicts the numeric values of two key dimensionless numbers and the allometry of the average reproductive allocation versus maturation size across species. Here we use this new model to predict the relationships of age-at-maturity, adult mortality and reproductive effort to environmental temperature and maturation size across species. Age-at-maturity, adult mortality and the proportion of the body mass given to reproduction per year are predicted to show ±0.25 power allometries with mass at maturity, and an exponential (Boltzmann) temperature dependence. Temperature is assumed to affect only body size growth, so the temperature linkages of maturation, mortality and reproductive effort are indirect via life history optimization; this is briefly contrasted with the idea that (for example) temperature directly affects mortality.
The majority of ectotherms grow slower but mature at a larger body size in colder environments. This phenomenon has puzzled biologists because classic theories of life-history evolution predict smaller sizes at maturity in environments that retard growth. During the last decade, intensive theoretical and empirical research has generated some plausible explanations based on nonadaptive or adaptive plasticity. Nonadaptive plasticity of body size is hypothesized to result from thermal constraints on cellular growth that cause smaller cells at higher temperatures, but the generality of this theory is poorly supported. Adaptive plasticity is hypothesized to result from greater benefits or lesser costs of delayed maturation in colder environments. These theories seem to apply well to some species but not others. Thus, no single theory has been able to explain the generality of temperature-size relationships in ectotherms. We recommend a multivariate theory that focuses on the coevolution of thermal reaction norms for growth rate and size at maturity. Such a theory should incorporate functional constraints on thermal reaction norms, as well as the natural covariation between temperature and other environmental variables.