Size Clines Research Articles

Little evidence for Bergmann's rule body size clines in passerines along tropical elevational gradients

AbstractAimTo test whether intra‐ and interspecific patterns in body mass along elevational gradients follow Bergmann's rule for a subset of tropical montane passerines.LocationTropical elevational gradients in New Guinea, Borneo, Peru and Costa Rica.MethodsI used linear regressions to assess intraspecific patterns in body mass along elevational gradients in common New Guinean passerines (2697 mist‐netted individuals of 21 species). I then evaluated interspecific patterns using two data sets. First, I investigated differences in body mass in species pairs of elevational replacements, closely related species with minimal overlap along elevational gradients (species pairs; New Guinea: n = 45, Borneo: n = 22, Peru: n = 58 and Costa Rica: n = 30). Second, I used a comparative phylogenetic approach to test whether species’ mid‐point elevations predicted their masses within entire passerine avifaunas found along single elevational gradients (species; New Guinea: n = 184, Peru: n = 529 and Costa Rica: n = 220).ResultsNew Guinean passerines exhibited minimal intraspecific variation in mass along elevational gradients. In two species, lower elevation individuals had significantly larger masses than upper elevation conspecifics. In species pairs of elevational replacements, there was no trend for upper elevation species to have larger masses than lower elevation species. Overall, species pairs tended to have positive mass disparities (mass of upper elevation species/mass of lower elevation species). However, contrary to predictions of Bergmann's rule, mass disparity was unrelated to elevational overlap. When considering entire passerine avifaunas along single elevational gradients, species’ masses were uncorrelated with their mid‐point elevational distributions.Main conclusionsI found little evidence that tropical montane passerines have larger body masses at higher elevations where temperatures are colder. This lack of pattern was consistent across evolutionarily independent avifaunas of different biogeographical regions. These results suggest mean temperature is not a generally important driver of body size evolution in tropical montane passerines.

Bergmann's rule in the oceans? Temperature strongly correlates with global interspecific patterns of body size in marine mammals

AbstractAimBergmann's rule remains unexplored in marine mammals. We first examine at a global extent whether these organisms show the same interspecific pattern reported for terrestrial mammals and then evaluate the influence of current environmental conditions and human impacts on the observed patterns.LocationGlobal.MethodsWe used range maps to document interspecific body size gradients and examined six environmental and human‐based hypotheses. We analysed the data using a comparative cross‐species method and a spatially explicit assemblage approach at three different grain sizes (200 km × 200 km, 400 km × 400 km and 800 km × 800 km). The associations between hypothesis‐linked predictors and body size were analysed through simple and multiple regressions that controlled for both spatial and phylogenetic autocorrelation.ResultsWe detected clear global latitudinal body size gradients, following a Bergmannian pattern (i.e. increasing size polewards). Consistently across methodological approaches (cross‐species and assemblage analyses) and grain sizes, sea surface temperature is the best predictor. Spatially, the temperature–size relationship is stronger in the Southern than in the Northern Hemisphere. Pinniped body sizes are critically constrained by temperature world‐wide whereas cetacean size clines show a weaker, albeit dominant, association with temperature.Main conclusionsAs in terrestrial mammals, our findings show that ambient temperature better explains interspecific body size patterns in cetaceans, and especially pinnipeds, world‐wide. Large‐bodied species are favoured in colder environments, in accordance with Bergmann's rule and the heat conservation hypothesis. However, our analyses also reveal a relevant role for salinity and primary productivity in migratory cetacean species. The large body sizes of baleen whales are essential for migration, for survival during fasting periods and minimizing the effects of temperature variation. This finding highlights the importance of spatially and phylogenetically explicit deconstructive approaches, considering alternative hypotheses to the traditional physiological mechanism, to gain a better understanding of Bergmann's rule.

Reevaluating geographic variation in life‐history traits of a widespread Nearctic amphibian

AbstractAnimals from cold environments are usually larger than animals from warm environments, which often produce clines in body size. Because variation in body size can lead to trade‐offs between growth and reproduction, life‐history traits should also vary across climatic gradients. To determine if life‐history traits of wood frogs Rana sylvatica vary with climate, we examined female and male body length, clutch size, and ovum size from 37 locations across an unprecedented 32° of latitude. In conflict with recent research, body size, and ovum size decreased in cold climates and at higher latitudes. Clutch size did not vary with climate or latitude, but reproductive effort (clutch size:female size) did, suggesting selection for a life‐history traits that favors maximizing propagule number over propagule size in cold climates. With accelerating climate change that will expose populations to novel environmental conditions, it is important to identify the limits of adaptation, which can be informed by greater understanding of variation in life‐history traits.

A large‐scale latitudinal pattern of life‐history traits in a strictly univoltine damselfly

1. Variation in thermal conditions and season length along latitudinal gradients affect body size‐related traits over different life stages. Selection is expected to optimise these size traits in response to the costs and benefits.2. Egg, hatchling, larval and adult size in males and females were estimated along a latitudinal gradient of 2730 km across Europe in the univoltine damselfly Lestes sponsa, using a combination of field‐collection and laboratory‐rearing experiments. In the laboratory, individuals were grown in temperatures and photoperiod simulating those at the latitude of origin, and in common‐garden conditions.3. The size of adults sampled in nature was negatively correlated with latitude. In all populations the females were larger than the males. Results from simulated and common‐garden rearing experiments supported this pattern of size difference across latitudes and between sexes, suggesting a genetic component for the latitudinal size trend and female‐biased size dimorphism. In contrast, hatchling size showed a positive relationship with latitude, but egg size, although differing between latitudes, showed no such relationship.4. The results support a converse Bergmann cline, i.e. a negative body size cline towards the north. This negative cline in body size is probably driven by progressively stronger seasonal time and temperature constraints towards the higher latitudes and by the obligate univoltine life cycle of L. sponsa. As egg size showed no relationship with latitude, other environmental factors besides temperature, such as desiccation risk, probably affect this trait.

Tests of ecogeographical relationships in a non-native species: what rules avian morphology?

The capacity of non-native species to undergo rapid adaptive change provides opportunities to research contemporary evolution through natural experiments. This capacity is particularly true when considering ecogeographical rules, to which non-native species have been shown to conform within relatively short periods of time. Ecogeographical rules explain predictable spatial patterns of morphology, physiology, life history and behaviour. We tested whether Australian populations of non-native starling, Sturnus vulgaris, introduced to the country approximately 150years ago, exhibited predicted environmental clines in body size, appendage size and heart size (Bergmann's, Allen's and Hesse's rules, respectively). Adult starlings (n=411) were collected from 28 localities from across eastern Australia from 2011 to 2012. Linear models were constructed to examine the relationships between morphology and local environment. Patterns of variation in body mass and bill surface area were consistent with Bergmann's and Allen's rules, respectively (small body size and larger bill size in warmer climates), with maximum summer temperature being a strongly weighted predictor of both variables. In the only intraspecific test of Hesse's rule in birds to date, we found no evidence to support the idea that relative heart size will be larger in individuals which live in colder climates. Our study does provide evidence that maximum temperature is a strong driver of morphological adaptation for starlings in Australia. The changes in morphology presented here demonstrate the potential for avian species to make rapid adaptive changes in relation to a changing climate to ameliorate the effects of heat stress.

Lifestyle-Based Approaches Provide Insights into Body Size Variation Across Environmental Gradients in Anurans

Body size of organisms as a fitness-related phenotype has evolved in response to local conditions, often through the size-dependent thermoregulatory mechanisms. The direction and degree of this response should depend on animals’ lifestyle in terms of the preference for terrestrial or aquatic conditions, especially so for adult anurans that differ in lifestyle among species but all must maintain certain body temperatures for metabolism. It may be expected that anuran species frequently exposed to terrestrial environments characterized by fluctuant thermal conditions are more plastic in body size along thermal gradients than those highly relaying on aquatic environments where thermal conditions are relatively stable. We test this prediction using both interspecific and intraspecific data. With anurans in China as the model organisms, we show that across terrestrial species but not aquatic species, body size decreases with increasing ambient temperature. From the published literature worldwide, we summarized that more terrestrial versus fewer aquatic species follow the predicted ecogeographical size patterns. In addition, both interspecific and intraspecific data reveal that arboreal anurans do not exhibit the size cline, probably because relatively warm climates experienced by these species impose weak selective pressures on heat conservation or adaptation to tree-climbing constrains the variation in body size. Our finding highlights the importance of taking lifestyle into account when assessing macroevolutionary trends in body size for anurans in particular and ectothermic taxa in general.

Body Size Adaptations to Altitudinal Climatic Variation in Neotropical Grasshoppers of the Genus Sphenarium (Orthoptera: Pyrgomorphidae).

Altitudinal clines in body size can result from the effects of natural and sexual selection on growth rates and developing times in seasonal environments. Short growing and reproductive seasons constrain the body size that adults can attain and their reproductive success. Little is known about the effects of altitudinal climatic variation on the diversification of Neotropical insects. In central Mexico, in addition to altitude, highly heterogeneous topography generates diverse climates that can occur even at the same latitude. Altitudinal variation and heterogeneous topography open an opportunity to test the relative impact of climatic variation on body size adaptations. In this study, we investigated the relationship between altitudinal climatic variation and body size, and the divergence rates of sexual size dimorphism (SSD) in Neotropical grasshoppers of the genus Sphenarium using a phylogenetic comparative approach. In order to distinguish the relative impact of natural and sexual selection on the diversification of the group, we also tracked the altitudinal distribution of the species and trends of both body size and SSD on the phylogeny of Sphenarium. The correlative evidence suggests no relationship between altitude and body size. However, larger species were associated with places having a warmer winter season in which the temporal window for development and reproduction can be longer. Nonetheless, the largest species were also associated with highly seasonal environments. Moreover, large body size and high levels of SSD have evolved independently several times throughout the history of the group and male body size has experienced a greater evolutionary divergence than females. These lines of evidence suggest that natural selection, associated with seasonality and sexual selection, on maturation time and body size could have enhanced the diversification of this insect group.

Wing trait-inversion associations in Drosophila subobscura can be generalized within continents, but may change through time.

Clinal variation is one of the most emblematic examples of the action of natural selection at a wide geographical range. In Drosophila subobscura, parallel clines in body size and inversions, but not in wing shape, were found in Europe and South and North America. Previous work has shown that a bottleneck effect might be largely responsible for differences in wing trait-inversion association between one European and one South American population. One question still unaddressed is whether the associations found before are present across other populations of the European and South American clines. Another open question is whether evolutionary dynamics in a new environment can lead to relevant changes in wing traits-inversion association. To analyse geographical variation in these associations, we characterized three recently laboratory founded D. subobscura populations from both the European and South American latitudinal clines. To address temporal variation, we also characterized the association at a later generation in the European populations. We found that wing size and shape associations can be generalized across populations of the same continent, but may change through time for wing size. The observed temporal changes are probably due to changes in the genetic content of inversions, derived from adaptation to the new, laboratory environment. Finally, we show that it is not possible to predict clinal variation from intrapopulation associations. All in all this suggests that, at least in the present, wing traits-inversion associations are not responsible for the maintenance of the latitudinal clines in wing shape and size.

Plastic Responses to Temperature Versus Local Adaptation at the Cold Extreme of the Climate Gradient

Climate is a strong selection agent at high elevations, but experimental examinations of how animals exclusive of highlands cope with its variation are scarce. We analysed temperature-induced variation of early ontogenetic traits in the alpine grasshopper Chorthippus cazurroi, and compared populations from the elevational extremes of the species distribution under laboratory conditions spanning natural temperature ranges. Neither elevation of origin, nor different growing temperatures, had a direct effect on nymph body size, but both factors contributed to size at hatching indirectly, via their effect on the duration of embryo development. Large emerging nymphs had a consistently greater survival, although small and fast-developing nymphs from highlands also performed well at low temperatures. Viability selection favoured fast-developing phenotypes in conditions in which plasticity delayed development, in a typical countergradient pattern. Growth in the successive stage did not compensate for slow development at hatching, thus responses at this early stage have potential long-lasting consequences. Although phenotypic selection during early development certifies the strength of selection imposed by cold temperatures in the laboratory, elevation clines of body size did not emerge in either nymphs or the wild parental generation. Differentiation in the wild may be levelled out by fecundity selection for large sizes, drift and gene flow resulting from the fragmentation and proximity of populations, or by micro-climatic differences that reduce the likelihood of directional selection. There is therefore potential for local adaptation to temperature, but a series of conditions typical of alpine environments and ectotherms may impair, confound or constrain full differentiation along the gradient.

Geographic body size variation in the periodical cicadas Magicicada: implications for life cycle divergence and local adaptation.

Seven species in three species groups (Decim, Cassini and Decula) of periodical cicadas (Magicicada) occupy a wide latitudinal range in the eastern United States. To clarify how adult body size, a key trait affecting fitness, varies geographically with climate conditions and life cycle, we analysed the relationships of population mean head width to geographic variables (latitude, longitude, altitude), habitat annual mean temperature (AMT), life cycle and species differences. Within species, body size was larger in females than males and decreased with increasing latitude (and decreasing habitat AMT), following the converse Bergmann's rule. For the pair of recently diverged 13- and 17-year species in each group, 13-year cicadas were equal in size or slightly smaller on average than their 17-year counterparts despite their shorter developmental time. This fact suggests that, under the same climatic conditions, 17-year cicadas have lowered growth rates compared to their 13-years counterparts, allowing 13-year cicadas with faster growth rates to achieve body sizes equivalent to those of their 17-year counterparts at the same locations. However, in the Decim group, which includes two 13-year species, the more southerly, anciently diverged 13-year species (Magicicada tredecim) was characterized by a larger body size than the other, more northerly 13- and 17-year species, suggesting that local adaptation in warmer habitats may ultimately lead to evolution of larger body sizes. Our results demonstrate how geographic clines in body size may be maintained in sister species possessing different life cycles.

An elevational trend of body size variation in a cold-climate agamid lizard, Phrynocephalus theobaldi

Abstract The pattern that many ectotherms have smaller body sizes in cold environments follows the converse to Bergmann’s rule and is most frequently found in lizards. Allen’s rule predicts animals from warm climates usually have longer tails and limbs, while these traits tend to be shorter in individuals from cold climates. We examined body size variation in an endemic Chinese lizard Phrynocephalus theobaldi along a broad elevational gradient (3,600–5,000 m on the Qinghai-Tibetan Plateau). Female body size showed a U-shaped cline, decreasing with increased elevation within the range 3,600–4,200 m, but increasing at elevations > 4200 m. Male body size continued to increase with increasing elevations. Both sexes showed an increased pattern of extremity length with elevation that does not conform to Allen’s rule. Limb length and tail length increased along the elevational gradients. In terms of color pattern, an abdominal black speckled area appears at elevations >4,200 m. This trait increases in size with increased elevation. Unlike most studies, our results indicated that annual sunshine hours corresponding to the activity period of the lizards could play an important role on the positive body size cline in environments at very high elevations > 4200 m.

Rethinking biogeographic patterns: high local variation in relation to latitudinal clines for a widely distributed species.

Wide-ranging species typically differ morphologically across their ranges. Bergmann's rule suggests that taxa in colder environments are bigger than related taxa in warmer locations. We examined 767 painted turtles (Chrysemys picta) in ten populations near their northwestern range edge in south-central British Columbia, Canada, in conjunction with previous data, to test the hypotheses of (1) a Bergmann's latitudinal cline, and (2) that males and females show similar latitudinal variation in size. We also explicitly test the impact of high local variation on range-wide inference. Female and male turtles showed similar latitudinal clines in body size; the degree of sexual dimorphism did not change across the range. Importantly, local variation in sexual dimorphism across ponds was nearly as high as the previously observed continental variation. Indeed, we found both the lowest and the highest degrees of sexual size dimorphism that have ever been reported for this species. Further, differing criteria in the literature for identifying mature females compound the difficulty of interpreting latitudinal clines in size or dimorphism. Our results highlight the need for much more systematic local and regional sampling as inputs for latitudinal or other comparative analyses such as Rensch's rule because insufficient sampling of high local variation may mask important ecological and evolutionary patterns.

Evidence for Bergmann's Rule and Not Allopatric Subspeciation in the Threatened Kaka (Nestor meridionalis).

Species of conservation concern characterized by small and declining populations greatly benefit from proactive management approaches such as population translocations. Because they often show intra-specific genetic and phenotypic variation, which can result from drift or differential selective pressures between habitats, understanding the distribution of such variation and its underlying processes is a prerequisite to develop effective management guidelines. Indeed, translocations among genetically differentiated populations potentially locally adapted are discouraged in order to avoid outbreeding depression, while translocations among populations characterized by high gene flow with no evidence for local adaptation are encouraged. Here, we first test whether 2 recognized subspecies, the North Island kaka (Nestor meridionalis septentrionalis) and South Island kaka (Nestor meridionalis meridionalis) of New Zealand fit a scenario of allopatric subspeciation following the separation of the North and South Islands at the end of the Pleistocene using 1 mtDNA (n = 96) and 9 microsatellite markers (n = 126). We then test whether morphological differences among the 2 subspecies support a pattern of local adaptation, comparing phenotypic divergence (P ST) and the level of divergence by drift alone (F ST) among populations. We find little population structure between islands, ruling out allopatric subspeciation in kaka. Further, P ST exceeds F ST, supporting an adaptive latitudinal size cline consistent with Bergmann's rule. These results therefore suggest that using neutral genetic diversity alone can be misleading when identifying management units and that the nature of phenotypic variation should be considered in translocations efforts. We finally discuss North and South Island management units but suggest that cross-island translocation be allowed.

Dispersal potential impacts size clines of grasshoppers across an elevation gradient

Body size is a life history trait that determines the reproductive success of a variety of organisms. Changes in body size may have a genetic component when persistent conditions such as season length and climate select for individuals of an optimal body size and an environmental component when it is influenced on an ecological scale by factors such as weather, food availability, or maternal effects. Along elevational gradients that experience seasonality, insects commonly become smaller with increases in elevation. In this study we test the hypothesis that dispersal potential, an indicator of gene flow, impacts the type of size clines exhibited by insects along elevational gradients and that these differences in local adaptation should lead to predictable changes in their reproductive potential and output. Using two short winged grasshopper species, Aeropedellus clavatus and Melanoplus boulderensis, and two long winged species, Camnula pellucida and Melanoplus sanguinipes, we showed that species with low dispersal potential are associated with significant declines in body size with increases in elevation while species with high dispersal potential displayed no size clines. Consistent with short winged species being more locally adapted, we show that reproductive potential, as measured by the proportion of ovarioles that become functional, do not differ among populations of short winged species, but decline with elevation in the long winged species. While our study failed to show that dispersal potential impacts reproductive output in a consistent and predictable manner (as measured by clutch and egg sizes), we address the possibility that clutch size may not reflect changes in total reproductive output and that changes in egg size may be a plastic trait. We concluded that studies exploring the evolution of body size, the reproductive capacity and species level responses to environmental change should note the importance of dispersal potential in influencing these patterns.

Environmental correlates of body size distribution in Cyprinidae (Actinopterygians) depend on phylogenetic scale

AbstractThe pattern of increasing species body size with increasing latitude has been noticed in different groups of animals. Here, we used seven key environmental factors and independent contrasts to assess body size latitudinal clines in Cyprinidae at two phylogenetic levels (inter‐ and intragenera), which were defined using a genus‐level supertree. Model selection procedures revealed that environmental factors shaping body size variation in Cyprinidae differed according to the phylogenetic scale considered. At the higher phylogenetic level, we found that both temperature (negative effect) and habitat availability (positive effect of drainage basin surface area) constituted mechanistic explanations of large‐scale body size distribution. No temperature‐related body size cline was observed at the intragenus level. Instead, competitive interaction (negative effect of species richness), habitat availability (positive effect of drainage basin surface area), migration ability and available energy (positive effects of glacial coverage and actual evapotranspiration) constitute alternative explanations at this lower phylogenetic scale. We conclude that (i) at the intergenus level, cyprinids do show a tendency to be smaller at high temperatures and larger at low temperatures, (ii) this tendency no longer exists at the intragenus level, (iii) latitude per se is a weak predictor of body size clines whatever the taxonomic level analysed, (iv) generalising geographical body size patterns may be rendered difficult by the superimposition of a series of mechanisms across different taxonomic scales, and (v) habitat size, here acting positively at both taxonomic scales, may play a major role in shaping riverine species body size clines.

Untangling intra- and interspecific effects on body size clines reveals divergent processes structuring convergent patterns in Anolis lizards.

Bergmann's rule-the tendency for body size to increase in colder environments-remains controversial today, despite 150 years of research. Considerable debate has revolved around whether the rule applies within or among species. However, this debate has generally not considered that clade-level relationships are caused by both intra- and interspecific effects. In this article, we implement a novel approach that allows for the separation of intra- and interspecific components of trait-environment relationships. We apply this approach to body size clines in two Caribbean clades of Anolis lizards and discover that their similar body size gradients are constructed in very different ways. We find inverse Bergmann's clines-high-elevation lizards are smaller bodied-for both the cybotes clade on Hispaniola and the sagrei clade on Cuba. However, on Hispaniola, the inverse cline is driven by interspecific differences, whereas intraspecific variation is responsible for the inverse cline on Cuba. Our results suggest that similar body size clines can be constructed through differing evolutionary and ecological processes, namely, through local adaptation or phenotypic plasticity (intraspecific clines) and/or size-ordered spatial sorting (interspecific clines). We propose that our approach can help integrate a divided research program by focusing on how the combined effects of intra- and interspecific processes can enhance or erode clade-level relationships at large biogeographic scales.

A comparative study of shell variation in two morphotypes of Lymnaea stagnalis (Mollusca: Gastropoda: Pulmonata)

Background Lymnaea stagnalis (L., 1758), the great pond snail, is among the most common and ubiquitous species of aquatic pulmonate gastropods of Palearctic. It is notorious for its enormous conchological variation, and many students tried to separate varieties, subspecies and even distinct species within the L. stagnalis s. l. Recent molecular studies have revealed that there are at least two genetically indistinguishable morphotypes of L. stagnalis in Palearctic that earlier were accepted by some authors as biological species under the binomial names L. stagnalis s. str. and L. fragilis (L., 1758). In this article, the comparative analysis of their conchological variation in a large physical geographical region (Western Siberia, Asiatic Russia) is provided.ResultsThe two morphotypes proved to be rather similar in their ecological preferences, but the patterns of their ontogenic, ecological, and geographical variation look rather distinct as well as the areas of their distribution in Western Siberia. U-shaped body size clines are reported in both morphotypes with the largest individuals tending to occur in the middle (forest-steppe) belt of Western Siberia. The causal analysis of the patterns of geographical variation in conchological traits of the great pond snail has identified the annual precipitation and the length of the growth season as the two main factors to shape the spatial clines in shell size and proportions. Among hydrological parameters, the water flow characteristics (lotic vs. lentic habitats) may influence shell morphology in the great pond snails though ecologically induced variation proved to be rather weak. The differences between populations living under different hydrological regimes may be captured by statistical techniques but are not enough to warrant separation of ‘ecological’ subspecies or other subspecific categories of ecophenotypic origin.ConclusionsThough there is no doubtless evidence of their specific independence, the two morphotypes, in a sense, ‘behave’ as two distinct entities with no identical ranges worthwhile to be recognized taxonomically. Their proper position in practical taxonomy should be discussed elsewhere.

Relative heart size in two rodent species increases with elevation: reviving Hesse's rule

AbstractAimWe assessed the phenotypic responses of body size and organ weights of two small rodent species (Myodes glareolus and Apodemus flavicollis) to elevation, a surrogate for temperature and other environmental conditions. We expected not only an increase in body size (Bergmann's rule), but also an increase in relative heart weight with increasing elevation (Hesse's rule).LocationThe Bohemian Forest, south‐eastern Germany.MethodsWe measured body length, body mass, and mass of the heart, lung, liver, kidneys and spleen of 386 adult specimens (188 of A flavicollis and 198 of M. glareolus) from 28 localities at 300–1450 m a.s.l., and also recorded the number of ectoparasites. We analysed body length using linear mixed models, with site as a random factor and species, sex, population density and elevation as fixed factors, and included all two‐way interactions between species and the remaining fixed variables. For the organ masses, we included body length and body mass in the calculations to account for allometric variation of organ mass with body size. For a subset of individuals, we sequenced the mitochondrial D‐loop to analyse genetic variation along the elevational gradient.ResultsFor both species, we found no support for Bergmann's rule. The body size of A. flavicollis even decreased with elevation. In contrast, the relative heart weight of both species significantly increased with elevation, thereby supporting Hesse's rule. Lung mass also increased with elevation. The mass of other internal organs showed no such consistent relationship with elevation. Neither species showed genetic differentiation across the elevational gradient.Main conclusionsOur results suggest that both rodent species respond to increasing elevation by decreasing overall energy expenditure and by increasing overall aerobic capacity. In the context of climate change, our results indicate that biogeographers should pay more attention to the potential significance of genetic and/or phenotypic plasticity of relative organ mass to environmental heterogeneity.

On the applicability of Bergmann’s rule to ectotherms: The state of the art

Bergmann’s rule (BR) is the classic ecogeographic principle that relates the body size of endotherms with environmental temperature (or latitude). The currently available data demonstrate that the latitudinal clines in body size predicted by BR are also observed in ectotherms, being fairly common in some groups. Despite plenty of published data, it is still impossible to estimate the frequency of Bergmann’s clines in ectotherms as thoroughly as has been done for endotherms. Within large taxa of ectotherms (Arthropoda, Mollusca, and Pisces), Bergmann’s clines occur along with converse Bergmann’s and U-shaped clines. Since the classic explanation of BR based on thermoregulation principles is not applicable to most ectotherms, quite a few hypotheses have been proposed that appeal to other foundations in search of such an explanation. Part of them suggests a direct modifying influence of temperature; however, most authors look for an adaptive sense of the increase in the body size of ectotherms at high latitudes and/or low temperatures. It appears that a single universal explanation of all the cases of Bergmann’s variation in ectotherms cannot be formulated. Most likely, the observed clines arise as a result of synergetic interactions between several factors covarying with latitude (or altitude above sea level). It is not always possible to estimate the exact contribution of each of these factors.

Ecogeographical and phylogenetic effects on craniofacial variation in macaques

The widespread and complex ecogeographical diversity of macaques may have caused adaptive morphological convergence among four phylogenetic subgroups, making their phylogenetic relationships unclear. We used geometric morphometrics and multivariate analyses to test the null hypothesis that craniofacial morphology does not vary with ecogeographical and phylogenetic factors. As predicted by Bergmann's rule, size was larger for the fascicularis and sinica groups in colder environments. No clear size cline was observed in the silenus and sylvanus groups. An allometric pattern was observed across macaques, indicating that as size increases, rounded faces become more elongated. However, the elevation was differentiated within each of the former two groups and between the silenus and sylvanus groups, and the slope decreased in each of the two northern species of the fascicularis group. All allometric changes resulted in the similar situation of the face being more rounded in animals inhabiting colder zones and/or in animals having a larger body size than that predicted from the overarching allometric pattern. For non-allometric components, variations in prognathism were significantly correlated with dietary differences; variations in localized shape components in zygomatics and muzzles were significantly correlated with phylogenetic differences among the subgroups. The common allometric pattern was probably influenced directly or indirectly by climate-related factors, which are pressures favoring a more rounded face in colder environments and/or a more elongated face in warmer environments. Allometric dissociation could have occurred several times in Macaca even within a subgroup because of their wide latitudinal distributions, critically impairing the taxonomic utility of craniofacial elongation.