Complex latitudinal variation in the morphology of the kleptoparasitic spider Argyrodes kumadai associated with host use and climatic conditions

We examined complex geographical patterns in the morphology of a kleptoparasitic spider, Argyrodes kumadai, across its distributional range in Japan. To disentangle biotic and abiotic factors underlying morphological variation, latitudinal trends were investigated in two traits, body size and relative leg length, across separate transition zones for host use and voltinism. Statistical analyses revealed complex sawtooth clines. Adult body size dramatically changed at the transition zones for host use and voltinism, and exhibited a latitudinal decline following the converse to Bergmann’s cline under the same host use and voltinism in both sexes. A similar pattern was observed for relative leg length in females but not in males. A genetic basis for a part of observed differences in morphology was supported by a common-garden experiment. Our data suggest that local adaptation to factors other than season length such as resource availability (here associated with host use) obscures underlying responses to latitude.     

Latitudinal insect body size clines revisited: a critical evaluation of the saw-tooth model

1.?Insect body size is predicted to increase with decreasing latitude because time available for growth increases. In insects with changing voltinism (i.e. number of generations per season), sharp decreases in development time and body size are expected at season lengths where new generations are added to the phenology of a species, giving rise to saw-tooth clines in these traits across latitudes. Growth rate variation may affect the magnitude of variation in body size or even reverse the saw-tooth cline. 2.?In this study, we analyse latitudinal body size clines in four geometrid moths with changing voltinism in a common laboratory environment. In addition to body size, we measured larval development time and growth rate and genetic correlations among the three traits. 3.?The patterns of clinal variation in body size were diverse, and the theory was not supported even when saw-tooth body size clines were found. Larval development time increased and growth rate decreased consistently with increasing season length, the clines in these traits being uniform. 4.?The consistencies of development time and growth rate clines suggest a common mechanism underlying the observations. Such a mechanism is discussed in relation to the complex interdependencies among the traits.     

Reaction norms for age and size at maturity in response to temperature: a test of the compound interest hypothesis

In ectotherms, temperature induces similar developmental and evolutionary responses in body size, with larger individuals occurring or evolving in low temperature environments. Based on the occasional occurrence of opposite size clines, showing a decline in body size with increasing latitude, an interaction between generation time and growing season length was suggested to account for the patterns found. Accordingly, multivoltine species with short generation times should gain high compound interest benefits from reproducing early at high temperatures, indicating potential for extra generations, even at the expense of being smaller. This should not apply for obligatorily monovoltine populations. We explicitly test the prediction that monovoltine populations (no compound interest) should be selected for large body size to maximise adult fitness, and therefore size at maturity should respond only weakly to temperature. In two monovoltine populations (an Alpine and a Western German one) of the butterfly Lycaena hippothoe, increasing temperatures had no significant effect on pupal weight and caused a slight decrease in adult weight only. In contrast, two closely related, yet potentially multivoltine Lycaena populations showed a greater weight loss at increasing temperature (in protandrous males, but not in females) and smaller adult sizes throughout. Thus, the results do support our predictions indicating that the compound interest hypothesis may yield causal explanations for the relationship between temperature and insect size at maturity. At all temperatures, the alpine population had higher growth rates and concomitantly shorter development times (not accompanied by a reduction in size) than the other, presumably indicating local adaptations to different climates.     

Season length,body size,and social polymorphism: size clines but not saw tooth clines in sweat bees

1. Annual insects are predicted to grow larger where the growing season is longer. However, transitions from one to two generations per year can occur when the season becomes sufficiently long, and are predicted to result in a sharp decrease in body size because available development time is halved. The potential for resulting saw‐tooth clines has been investigated only in solitary taxa with free‐living larvae. 2. Size clines were investigated in two socially polymorphic sweat bees (Halictidae): transitions between solitary and social nesting occur along gradients of increasing season length, characterised by the absence or presence of workers and offspring that are individually mass provisioned by adults. How the body size changes with season length was examined, and whether transitions in social phenotype generate saw‐tooth size clines. We measured Lasioglossum calceatum and Halictus rubicundus nest foundresses originating from more than 1000 km of latitude, encompassing the transition between social and solitary nesting. 3. Using satellite‐collected temperature data to estimate season length, it was shown that both species were largest where the season was longest. Body size increased linearly with season length in L. calceatum and non‐linearly in H. rubicundus but the existence of saw‐tooth clines was not supported. 4. The present results suggest that because the amount of food consumed by offspring during development is determined by adults, environmental and social influences on the provisioning strategies of adult bees may be more important factors than available feeding time in determining offspring body size in socially polymorphic sweat bees.     

Body size clines in the European badger and the abundant centre hypothesis

Aim To test the abundant centre hypothesis by analysing the physical and climatic factors that influence body size variation in the European badger (Meles meles). Location Data were compiled from 35 locations across Europe. Methods We used body mass, body length and condylo‐basal length (CBL) as surrogates of size. We also compiled data on latitude, several climatic variables, habitat type and site position relative to the range edge. We collapsed all continuous climatic variables into independent vectors using principal components analysis (PCA), and used a general linear model to explain the morphometric variation in badger populations across the species’ range. Results Body mass and body length were nonlinearly and significantly related to latitude. In contrast, CBL was linearly related to latitude. Body mass changed nonlinearly along the temperature (PC1) gradient, with the highest values observed at mid‐range. Furthermore, body mass, body length and CBL differed significantly among habitats, with badgers showing larger size in temperate habitats and core areas relative to peripheral zones. Main conclusions Our analysis supports the nonlinear pattern predicted by the abundant centre hypothesis only for body mass and body length. These results imply that individuals are largest and heaviest at the centre of the climatic range of badger distribution. Variation of CBL with latitude follows a linear trend, consistent with Bergmann’s rule. Our results provide mixed support for the abundant centre hypothesis, and suggest food availability/quality to be the main mechanism underlying body size clines in this species.     

Latitudinal pattern in body size in a cockroach,Eupolyphaga sinensis

Body size of insects with flexible life cycles is expected to conform to the saw‐tooth model, a model in which the relationship between size and developmental time depends on length of the growing season. In species with high variability in terms of voltinism, however, more complex patterns can be expected. Few empirical studies have demonstrated the existence of such relationships, or whether climatic factors contribute to these relationships. In this study, we investigated the geographic variation in body size of the Chinese cockroach, Eupolyphaga sinensis Walker (Blattaria: Polyphagidae), which has a variable life cycle length. The sizes of adults – collected from 14 localities ranging from temperate to subtropical zones in China – were measured, using body length, body width, and pronotum width as parameters. The relationship between size, latitude, and climate factors (encompassing 10 variables) was then investigated. We found that the body size of E. sinensis varied considerably with latitude: cockroaches were larger at low and high latitudes, but smaller at intermediate latitudes. Thus, the relationship between climate and body size conformed to a saw‐tooth pattern. Results indicate that two factors were significantly associated with body size clines: season length and variability in life cycle length. Our results also demonstrated that climatic factors contribute to latitudinal clines in body size, which has important ecological and evolutionary implications. It can be expected that global climate change may alter latitudinal clines in body size of E. sinensis.     

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.     

Habitat use, diet and body size of Heard Island weevils

Habitat use, diet and body-size variation are examined in weevils from Heard Island, with specific attention being given to the Ectemnorhinus viridis species complex. E. viridis shows marked altitudinal variation in body size and vestiture, but there are no consistent associations between body size and diet, nor are there consistent among-individual differences in conventional taxonomic characters. Thus, the status of E. viridis as a single, variable species is maintained. This species occurs from sea level to 600 m and it feeds on vascular plants and bryophytes. Canonopsis sericeus also feeds on bryophytes and vascular plants and occurs over a narrower altitudinal range. Palirhoeus eatoni is restricted to the surpralittoral zone where it feeds on marine algae and lichens. Bothrometopus brevis and B. gracilipes both feed on cryptogams, with the former species occurring from sea level to 450 m, and the latter from 50 to 550 m above sea level. In all species, males are smaller than females and there is a size cline such that populations from higher elevations are smaller than those at lower altitudes. This cline is the reverse of that found on the Prince Edward Islands which, unlike Heard Island, lie to the north of the Antarctic Polar Frontal Zone. This difference in body-size clines between weevils on the two island groups is ascribed to the shorter growing season on the colder Heard Island. The information presented here supports previous ideas regarding the evolution of the Ectemnorhinus-group of weevils on the South Indian Ocean Province Islands, although it suggests that subsequent tests of these hypotheses would profit from the inclusion of molecular systematic work.     

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.     

Countergradient variation in growth rate: compensation for length of the growing season among Atlantic silversides from different latitudes

Summary How do organisms adapt to the differences in temperature and length of the growing season that occur with latitude? Among Atlantic silversides (Menidia menidia) along the east coast of North America, the length of the first growing season declines by a factor of about 2.5 with increasing latitude. Yet body size at the end of the first growing season does not decline. High-latitude fish must, therefore, grow faster within the growing season than do low-latitude fish. This geographical pattern has a genetic basis. Laboratory experiments on fish from six different locations revealed a latitudinal gradient in the capacity for growth (i.e., maximum growth potential). In two subsequent experiments using fish from Nova Scotia (NS), New York (NY) and South Carolina (SC) that had been separately reared in a common environment for several generations, differences in growth rate among populations were highly significant. The rank order was NS>NY>SC, but the difference among populations depended on temperature. High-latitude fish outperformed those from low latitudes primarily at the high temperatures that low-latitude fish would be expected to experience most often in nature. These results suggest that instead of being adapted for growth at low temperatures, fish from high latitudes are adapted for rapid elevation of growth rate during the brief interval of the year when high temperatures occur. Selection on growth rate results from sizedependent winter mortality: the importance to winter survival of being large increases with latitude but the length of the growing season simultaneously decreases. The end result is countergradient variation in growth rate, a phenomenon that may be much more widespread than currently recognized.     

What determines conformity to Bergmann's rule?

Aim  Bergmann's rule, the tendency of body size within species in bird and mammal populations to be positively correlated with latitude, is among the best known biogeographical generalizations. The factors behind such clines, however, are not well understood. Here we use a large data base of 79 mammalian carnivore species to examine the factors affecting latitudinal size clines.
Location  Worldwide.
Methods  We measured the skulls and teeth of carnivores in natural history museums, and calculated the amount of variation in size explained by latitude, supplementing our measurements with published data. We examined the effects of a number of variables on the tendency to show latitudinal clines.
Results  We found that geographical range and latitudinal extent are strongly related to size clines. Minimum temperatures across the range, net primary productivity and habitat diversity also have some, albeit much less, influence.
Main conclusions  We suggest that species with large geographical ranges are likely to encounter significant heterogeneity in those factors that influence body size, and are thus likely to exhibit size clines. However, the key factors that determine body size may not always operate along a latitudinal (or other geographical) cline, but be spatially linked to patches in the species range. One such important factor is likely to be food availability, which we show is a strong predictor of size in the brown bear ( Ursus arctos ) but is not associated with a latitudinal cline. We argue that the spatial distribution of key resources within the species range constitutes a significant predictor of carnivore body size.     

Selection does not favor larger body size at lower temperature in a seed-feeding beetle

Body size of many animals increases with increasing latitude, a phenomenon known as Bergmann's rule (Bergmann clines). Latitudinal gradients in mean temperature are frequently assumed to be the underlying cause of this pattern because temperature covaries systematically with latitude, but whether and how temperature mediates selection on body size is unclear. To test the hypothesis that the "relative" advantage of being larger is greatest at cooler temperatures we compare the fitness of replicate lines of the seed beetle, Stator limbatus, for which body size was manipulated via artificial selection ("Large,"Control," and "Small" lines), when raised at low (22 degrees C) and high (34 degrees C) temperatures. Large-bodied beetles (Large lines) took the longest to develop but had the highest lifetime fecundity, and highest fitness (r(C)), at both low and high temperatures. However, the relative difference between the Large and Small lines did not change with temperature (replicate 2) or was greatest at high temperature (replicate 1), contrary to the prediction that the fitness advantage of being large relative to being small will decline with increasing temperature. Our results are consistent with two previous studies of this seed beetle, but inconsistent with prior studies that suggest that temperature-mediated selection on body size is a major contributor to the production of Bergmann clines. We conclude that other environmental and ecological variables that covary with latitude are more likely to produce the gradient in natural selection responsible for generating Bergmann clines.     

Bergmann and converse bergmann latitudinal clines in arthropods: two ends of a continuum?

Two seemingly opposite evolutionary patterns of clinal variationin 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. Athird pattern causally related to the latter is countergradientvariation, whereby populations of a given species compensateseasonal limitations at higher latitudes by evolving fastergrowth and larger body sizes compared to their low latitudeconspecifics. We discuss these patterns and argue that theyare not mutually exclusive because they are driven by differentenvironmental causes and proximate mechanisms; they thereforecan act in conjunction, resulting in any intermediate pattern.Alternatively, Bergmann and converse Bergmann clines can beinterpreted as over- and undercompensating countergradient variation,respectively. We illustrate this with data for the wide-spreadyellow dung fly, Scathophaga stercoraria (Diptera: Scathophagidae),which in Europe shows a Bergmann cline for size and a converseBergmann cline (i.e., countergradient variation) for developmenttime. A literature review of the available evidence on arthropodlatitudinal clines further shows a patterned continuum of responses.Converse Bergmann clines due to end-of-season time limitationsare more common in larger species with longer development times.Our study thus provides a synthesis to the controversy aboutthe importance of Bergmann's rule and the converse Bergmannrule in nature.     

Do southern elephant seals show density dependence in fecundity?

Here we provide an alternative interpretation to that of Pistorius et al. (2001), concerning density-dependent increases in fecundity resulting in population regulation of the southern elephant seal population at Marion Island. We do not contradict the findings of Pistorius et al. (2001), because it does appear: (1) that a change in fecundity has been observed, and (2) that some factor related to food supply is the most likely cause for an observed population decline and increase in reproductive performance. The main observation leading to the interpretation of density-dependent feedback in the population of southern elephant seals at Marion Island (one of the Prince Edward Islands) is that there has been a reduction in the population's rate of decline in recent years (reported by Pistorius et al. (1999b)), and that this could have resulted from a per capita increase in food availability. However, because rates of population change are rarely linearly constant, changes in population size should be expressed on a logarithmic, rather than a linear scale, as used by Pistorius et al. (1999b). Re-plotting the linear values of Pistorius et al. (1999b) on the natural logarithmic scale gave no clear change in the rate of population decline; therefore, we conclude that the rate of population change (decline) has remained constant from 1986 to 1997 (r=-0.0439). The Marion Island population is part of the larger Kerguelen population, and there might be considerable overlap in the foraging areas, and possibly prey, exploited by elephant seals from all sub-populations within this larger population. Changes in the number of intra-specific resource competitors at Marion Island are therefore unlikely to alter per capita food availability since the Marion population constitutes approximately 1% of the total Kerguelen population. We propose an alternative hypothesis that the present data support a mechanism driving the proposed increase in per capita food supply through changes in either: (1) inter-specific food competition, (2) rates of predation, (3) changes in weather pattern or (4) disease.     

Optimizing development time in a seasonal environment: The ‘ups and downs’ of clinal variation

Summary This paper explores the problem of adapting development time to changes in the length of time conditions are favourable for growth and reproduction (season length). It is shown that systematic changes in season length along some gradient such as latitude can generate either simple clines in development time or saw-tooth clines. The relationship between development time and body size gives rise to a corresponding variation in body size. The generation of a saw-tooth cline does not require sharp environmental changes. Both types of clinal variation are observed in insects.     

Testing mechanisms of Bergmann's rule: phenotypic decline but no genetic change in body size in three passerine bird populations

Bergmann's rule predicts a decrease in body size with increasing temperature and has much empirical support. Surprisingly, we know very little about whether "Bergmann size clines" are due to a genetic response or are a consequence of phenotypic plasticity. Here, we use data on body size (mass and tarsus length) from three long-term (1979-2008) study populations of great tits (Parus major) that experienced a temperature increase to examine mechanisms behind Bergmann's rule. We show that adult body mass decreased over the study period in all populations and that tarsus length increased in one population. Both body mass and tarsus length were heritable and under weak positive directional selection, predicting an increase, rather than a decrease, in body mass. There was no support for microevolutionary change, and thus the observed declines in body mass were likely a result of phenotypic plasticity. Interestingly, this plasticity was not in direct response to temperature changes but seemed to be due to changes in prey dynamics. Our results caution against interpreting recent phenotypic body size declines as adaptive evolutionary responses to temperature changes and highlight the importance of considering alternative environmental factors when testing size clines.     

Correlated Changes in Body Melanisation and Mating Traits of Drosophila melanogaster: A Seasonal Analysis

Altitudinal localities of the northern India are associated with high seasonal changes. Drosophila melanogaster flies are darker during the winter season as compared to the autumn season. We tested the hypothesis whether there are altitudinal clines for mating related traits. We observed negative cline for mating latency and positive for copulation period along altitude in D. melanogaster. We further tested if seasonally varying body melanisation is correlated with mating propensity in D. melanogaster. Thus, we examined the D. melanogaster flies collected during autumn and winter season for changes in body melanisation and mating-related traits. Flies from the winter season show high melanisation, copulation duration and fecundity/day as compared to the autumn season flies. By contrast mating latency is longer during autumn as compared to winter season. Based on within- and between-population analysis, body melanisation shows positive correlation with copulation duration and fecundity/day, while negative correlation with mating latency. Within-population analyses show no correlation between body size and ovariole number with body melanisation. Thus, our data suggest that seasonal changes in body melanisation are correlated with mating latency, copulation duration and fecundity/day, but no correlation with body size and ovariole numbers. Further, we observed that seasonal changes in these clines, although have some component of plasticity, have strong genetic basis as the seasonal and population differences were maintained for various traits after 8 generations in the laboratory.     

Local climate determines intra‐ and interspecific variation in sexual size dimorphism in mountain grasshopper communities

The climate is often evoked to explain broad‐scale clines of body size, yet its involvement in the processes that generate size inequality in the two sexes (sexual size dimorphism) remains elusive. Here, we analyse climatic clines of sexual size dimorphism along a wide elevation gradient (i) among grasshopper species in a phylogenetically controlled scenario and (ii) within species differing in distribution and cold tolerance, to highlight patterns generated at different time scales, mainly evolutionary (among species or higher taxa) and ontogenetic or microevolutionary (within species). At the interspecific level, grasshoppers were slightly smaller and less dimorphic at high elevations. These clines were associated with gradients of precipitation and sun exposure, which are likely indicators of other factors that directly exert selective pressures, such as resource availability and conditions for effective thermoregulation. Within species, we found a positive effect of temperature and a negative effect of elevation on body size, especially on condition‐dependent measures of body size (total body length rather than hind femur length) and in species inhabiting the highest elevations. In spite of a certain degree of species‐specific variation, females tended to adjust their body size more often than males, suggesting that body size in females can evolve faster among species and can be more plastic or dependent on nutritional conditions within species living in adverse climates. Natural selection on female body size may therefore prevail over sexual selection on male body size in alpine environments, and abiotic factors may trigger consistent phenotypic patterns across taxonomic scales.     

Humidity and seasonality drives body size patterns in males of the bush cricket Isophya rizeensis Sevgili, 2003 （Orthoptera： Tettigoniidae： Phaneropterinae）

Two primary patterns of body size variation have been recorded in ectotherms in relation to latitudinal/altitudinal shifts. In some, body size increases with increasing latitude/altitude whereas, in others, body size decreases with increasing latitude/altitude. This clinal variation is generally assumed to be caused by local adaptation to environmental conditions however the selective variable（s） （temperature, humidity, diet quality, etc.） is still heavily debated. Here we investigate geographic variation in body size of dark and pale color morphs of males of the bush-cricket lsophya rizeensis collected from 15 locations along an elevation gradient ranging from 350 to 2 500 m. Using an information theoretical approach we evaluate the relative support of four different hypotheses （the temperature size rule, the moisture gradient hypothesis, the seasonal constraint hypothesis, and the primary productivity hypothesis） explaining body size variation along the altitudinal gradient. Body size variation in pale color morphs showed a curvilinear relationship with altitude while dark color morphs showed no variation in body size. Body size variation in pale color morphs was highly correlated with precipitation and temperature seasonality values thus giving strong support for the moisture gradient and seasonal constraint hypothesis. Our results reinforce the importance of gradients in humidity and seasonality over temperature in the creation of altitudinal body size clines and the role of selection for resistance to stress factors in the establishment of these clines. Whether a body size cline is observed or not might also depend on the phenotypic properties of the individuals, like coloration.