GEOGRAPHIC VARIATION IN LIFE-HISTORY TRAITS OF THE ANT LION,MYRMELEON IMMACULATUS: EVOLUTIONARY IMPLICATIONS OF BERGMANN'S RULE

In eastern North America, body size of the larval ant lion Myrmeleon immaculatus increases from south to north, following Bergmann's rule. We used a common-garden experiment and a reciprocal-transplant experiment to evaluate the effects of food and temperature on ant lion growth, body size, and survivorship. In the laboratory common-garden experiment, first-instar larvae from two southern (Georgia, South Carolina) and two northern (Connecticut, Rhode Island) populations were reared in incubators under high- and low-food and high- and low-temperature regimes. For all populations, high food increased final body mass and growth rate and decreased development time. Growth rates were higher at low temperatures, but temperature did not affect larval or adult body mass. Survivorship was highest in high-food and low-temperature treatments. Across all food and temperature treatments, northern populations exhibited a larger final body mass, shorter development time, faster growth rate, and greater survivorship than did southern populations. Results were similar for a field reciprocal-transplant experiment of third-instar larvae between populations in Connecticut and Oklahoma: Connecticut larvae grew faster than Oklahoma larvae, regardless of transplant site. Conversely, larvae transplanted to Oklahoma grew faster than larvae transplanted to Connecticut, regardless of population source. These results suggest that variation in food availability, not temperature, may account for differences in growth and body size of northern and southern ant lions. Although northern larvae grew faster and reached a larger body size in both experiments, northern environments should suppress growth because of reduced food availability and a limited growing season. This study provides the first example of countergradient selection causing Bergmann's rule in an ectotherm.     

Test for latitudinal variation of life history,behavior and mortality in the strictly univoltine damselfly Sympecma fusca (Zygoptera: Lestidae)

A wealth of evidence shows that combinations of ecological stressors interact in shaping life history traits, but little is known about how ecological stressors combine with different seasonal time constraints to shape life history, behavior and mortality across populations. We studied life history, behavior and mortality rate in two latitudinally distant populations of the strictly univoltine, adult‐overwintering damselfly Sympecma fusca. Results from laboratory common‐garden and outdoor experiments indicated countergradient variation of larval development time and growth rate: the more time‐constrained larvae showed faster development and a higher growth rate. This finding led to larger size at emergence in the more time‐constrained individuals. Under conditions of intraspecific interaction (outdoor experiment), northern individuals showed lower survival than southern ones, presumably due to cannibalism. In the absence of intraspecific interactions (laboratory experiment), northern and southern larvae did not differ in survival. Finally, laboratory‐grown northern and southern larvae did not differ in activity level. This is the first time that compensation for seasonal time constraints has been shown in a temperate odonate species that overwinters in the adult stage.     

Are high-latitude individuals superior competitors? A test with <Emphasis Type="Italic">Rana temporaria</Emphasis> tadpoles

Species with a wide distribution over latitudinal gradients often exhibit increasing growth and development rates towards higher latitudes. Ecological theory predicts that these fast-growing genotypes are, in the absence of trade-offs with fast growth, better competitors than low-latitude conspecifics. While knowledge on key ecological traits along latitudinal clines is important for understanding how these clines are maintained, the relative competitive ability of high latitude individuals against low latitude conspecifics has not been tested. Growth and development rates of the common frog Rana temporaria increase along the latitudinal gradient across Scandinavia. Here we investigated larval competition over food resources within and between two R. temporaria populations originating from southern and northern Sweden in an outdoor common garden experiment. We used a factorial design, where southern and northern tadpoles were reared either as single populations or as mixes of the two populations at two densities and predator treatments (absence and non-lethal presence of Aeshna dragonfly larvae). Tadpoles from the high latitude population grew and developed faster and in the beginning of the experiment they hid less and were more active than tadpoles from the low latitude population. When raised together with high latitude tadpoles the southern tadpoles had a longer larval period, however, the response of high latitude tadpoles to the competition by low latitude tadpoles did not differ from their response to intra-population competition. This result was not significantly affected by density or predator treatments. Our results support the hypothesis that high latitude populations are better competitors than their low latitude conspecifics, and suggest that in R. temporaria fast growth and development trade off with other fitness components along the latitudinal gradient across Scandinavia.     

Two experimental designs generate contrasting patterns of behavioral differentiation along a latitudinal gradient in Lestes sponsa—Common‐garden not so common after all?

Understanding why and how behavioral profiles differ across latitudes can help predict behavioral responses to environmental change. The first response to environmental change that an organism exhibits is commonly a behavioral response. Change in one behavior usually results in shifts in other correlated behaviors, which may adaptively or maladaptively vary across environments and/or time. However, one important aspect that is often neglected when studying behavioral expressions among populations is if/how the experimental design might affect the results. This is unfortunate since animals often plastically modify their behavior to the environment, for example, rearing conditions. We studied behavioral traits and trait correlations in larvae of a univoltine damselfly, Lestes sponsa, along its latitudinal distribution, spreading over 3,300 km. We compared behavioral profiles among larvae grown in two conditions: (a) native temperatures and photoperiods or (b) averaged constant temperatures and photoperiods (common‐garden). We hypothesized latitudinal differences in behavioral traits regardless of the conditions in which larvae were grown, with northern populations expressing higher activity, boldness, and foraging efficiency. When grown in native conditions, northern larvae were bolder, more active and more effective in prey capture than central and low latitude populations, respectively, as well as showed the strongest behavioral correlations. In contrast, larvae reared in common‐garden conditions showed no differences between regions in both individual traits and trait correlations. The results suggest different selective pressures acting on the studied traits across populations, with environment as a central determinant of the observed trait values. Common‐garden designed experiments may evoke population‐dependent levels of plastic response to the artificial conditions and, hence, generate results that lack ecological relevance when studying multi‐population differences in behavior.     

Environmental and genetic effects on flowering differences between northern and southern populations of Arabidopsis lyrata (Brassicaceae)

Arabidopsis lyrata (Brassicaceae) is a close outcrossing relative of A. thaliana. We examine flowering time variation of northern and southern A. lyrata populations in controlled environmental conditions, in a common garden experiment with A. thaliana, and in the field. Southern populations of A. lyrata flowered earlier than northern ones in all environmental conditions. Individuals from southern populations were more likely to flower in short days (14 h light) than northern ones, and all populations had a higher probability of flowering and flowered more rapidly in long days (20 h). The interaction of population and day length significantly affected flowering probability, and flowering time in one of two comparisons. The common garden experiment demonstrated differences between populations in the response to seed cold treatment, but growth chamber experiments showed no vernalization effect after 4 wk of rosette cold treatment. In a field population in Norway, a high proportion of the plants flowered in each year of the study. The plants progressed to flowering more rapidly in the field and common garden than in the growth chamber. The genetic basis of these flowering time differences here can be further studied using A. thaliana genetic tools.     

Rapid divergence in physiological and life-history traits between northern and southern populations of the British introduced neo-species, Senecio squalidus

Alien plants provide a unique opportunity to study evolution in novel environments, but relatively little is known about the extent to which they become locally adapted to different environments across their new range. Here, we compare northern and southern populations of the introduced species Senecio squalidus in Britain; S. squalidus has been in southern Britain for approximately 200  years and reached Scotland only about 50  years ago. We conducted common garden experiments at sites in the north and south of the species' range in Britain. We also conducted glasshouse and growth chamber experiments to test the hypothesis that southern genotypes flower later, are more drought-tolerant, germinate and establish better at warmer temperatures, and are less sensitive to cold stress than their more northern counterparts. Results from the common garden experiments are largely consistent with the hypothesis of rapid adaptive divergence of populations of the species within the introduced range, with genotypes typically showing a home-site advantage. Results from the glasshouse and growth chamber experiments demonstrate adaptive divergence in ability to tolerate drought stress and high temperatures, as well as in phenology. In particular, southern genotypes were more tolerant of dry conditions and high temperatures and they flowered later than northern genotypes. Our results show that rapid local adaptation can occur in alien species, and they have implications for our understanding of the ecological genetics of range expansion of introduced weeds.     

Life‐history differences across latitude in common side‐blotched lizards (Uta stansburiana)

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An experimental study on the foraging behavior of a pit-building antlion larva,Myrmeleon bore

Foraging behavior of a pit-building antlion larva, Myrmeleon boreTjeder was investigated experimentally to elucidate the relation between the feeding level and pit relocation.

1. In artificial sands constructed in the field the 3rd instar larvae of M. bore rarely changed the positions of their pits, though several antlions had moved actively until they constructed pits. The average feeding rate was 0.3 prey/day/pit, and about 60% of prey captured were ants.
2. To examine whether or not M. bore larvae concentrate into the area where they can capture more prey, 8 antlions were released into each of 6 boxes filled with sand. I divided the sand surface of each box into two half areas, then gave prey to the pits built in a half area and gave no prey to the pits built in the other half. During the 50-day observation period, nonfed antlions never moved into the area where prey were given.
3. The 3rd instar larvae were reared separately without food. Even under starved conditions they rarely relocated their pits until dealth. The average duration of survival period was 83.9 days.
4. The experimental results indicate that M. bore larvae adopt a tactic of sedentary ambushing. These larvae exhibit low movement rates which are independent of prey capture rates.

     

Variation in the degree and costs of adaptive phenotypic plasticity among Rana temporaria populations

Adaptive phenotypic plasticity in the form of capacity to accelerate development as a response to pond drying risk is known from many amphibian species. However, very little is known about factors that might constrain the evolution of this type of plasticity, and few studies have explored to what degree plasticity might be constrained by trade-offs dictated by adaptation to different environmental conditions. We compared the ability of southern and northern Scandinavian common frog (Rana temporaria) larvae originating from 10 different populations to accelerate their development in response to simulated pond drying risk and the resulting costs in metamorphic size in a factorial laboratory experiment. We found that (i) northern larvae developed faster than the southern larvae in all treatments, (ii) a capacity to accelerate the response was present in all five southern and all five northern populations tested, but that the magnitude of the response was much larger (and less variable) in the southern than in the northern populations, and that (iii) significant plasticity costs in metamorphic size were present in the southern populations, the plastic genotypes having smaller metamorphic size in the absence of desiccation risk, but no evidence for plasticity costs was found in the northern populations. We suggest that the weaker response to pond drying risk in the northern populations is due to stronger selection on large metamorphic size as compared with southern populations. In other words, seasonal time constraints that have selected the northern larvae to be fast growing and developing, may also constrain their innate ability for adaptive phenotypic plasticity.     

Influence of seasonal time constraints on growth and development of common frog tadpoles: a photoperiod experiment

Organisms living in seasonal environments are often limited by the time available to complete their development. Especially individuals in northern populations may face severe time constraints in their need of completing development before the end of the growth season. Larval amphibians have been widely used in studies of phenotypic plasticity. However, their responses to changes in photoperiod, the main seasonal cue in many organisms, are unknown. In a laboratory experiment, we studied whether common frog (Rana temporaria) tadpoles originating from two populations (separated latitudinally by 1600 km) adjust their growth and development according to the progress of the season by using photoperiodic cues, and whether these responses are temperature dependent. We hypothesised that if frogs use photoperiod as a cue, they should increase growth and development rates as a response to photoperiodic treatments mimicking progressing season. Although our predictions were not verified in either of the populations, photoperiod manipulations had effects on larval life history in both populations. When exposed to progressing season treatments and high temperature, tadpoles from the southern population ceased feeding, which led to delayed metamorphosis and increased mortality. In the northern population, age at metamorphosis was unaffected by the photoperiod treatments, but growth rate until metamorphosis and metamorphic size were slightly larger in the treatments with shorter (increasing or decreasing) day length. Irrespective of photoperiod treatments, growth and development rates, size at metamorphosis and food consumption were higher in the northern as compared to the southern population. These results indicate that in contrast to several insect species, the critical life history decisions in amphibian larvae may not be strongly influenced by photoperiodic cues, but different populations seem to differ in this respect. However, the strong temperature×photoperiod interactions in several traits in the southern population suggest that the role of photoperiodic cues may be affected by other environmental factors, although the ecological significance of these differences remains unclear.     

Climatic adaptation in an isolated and genetically impoverished amphibian population

The capacity of populations to respond adaptively to environmental change is essential for their persistence. Isolated populations often harbour reduced genetic variation, which is predicted to decrease adaptive potential, and can be detrimental under the current scenarios of global change. In this study, we examined climatic adaptation in larval life history traits in the pool frog Rana lessonae along a latitudinal gradient across the northern distribution area of the species, paying special attention to the isolated and genetically impoverished fringe populations in central Sweden. Larvae from eight populations within three geographic areas (Poland, Latvia and Sweden) were reared under three temperatures (19, 22 and 26°C) in a common garden laboratory experiment. We found clear evidence for latitudinal adaptation in R. lessonae populations, larvae from higher latitudes growing and developing faster than low‐latitude ones. Larvae from the Swedish populations were able to compensate for the effects of cooler temperatures and a shorter growth season with genetically higher growth and development rates (i.e. countergradient variation) in the two higher temperature treatments, but there was no difference among the populations at the lowest temperature treatment, which is likely to be close to the temperature limiting growth in R. lessonae. Our results demonstrate that isolated and genetically impoverished populations can be locally adapted, and identify the Swedish fringe populations as a significant conservation unit adapted to the northern environmental conditions.     

Natural Stressors and Ranavirus Susceptibility in Larval Wood Frogs (Rana sylvatica)

Chronic exposure to stressors has been shown to suppress immune function in vertebrates, making them more susceptible to pathogens. It is less clear, however, whether many natural stressors are immunosuppressive. Moreover, whether stressors make disease more likely or more severe in populations is unclear because animals respond to stressors both behaviorally and physiologically. We tested whether chronic exposure to three natural stressors of wood frog tadpoles—high-densities, predator-cues, and low-food conditions—influence their susceptibility to a lethal ranavirus both individually in laboratory experiments, and collectively in outdoor mesocosms. Prior to virus exposure, we observed elevated corticosterone only in low-food treatments, although other treatments altered rates of growth and development as well as tadpole behavior. None of the treatments, however, increased susceptibility to ranavirus as measured by the proportion of tadpoles that became infected or died, or the time to death compared to controls. In fact, mortality in the mesocosms was actually lower in the high-density treatment even though most individuals became infected, largely because of increased rates of metamorphosis. Overall we find no support for the hypothesis that chronic exposure to common, ecologically relevant challenges necessarily elevates corticosterone levels in a population or leads to more severe ranaviral disease or epidemics. Conditions may, however, conspire to make ranavirus infection more common in metamorphosing amphibians.     

How caterpillars avoid overheating: behavioral and phenotypic plasticity of pipevine swallowtail larvae

We tested the hypothesis that larvae of the pipevine swallowtail butterfly, Battus philenor, employ behavioral and phenotypic plasticity as thermoregulatory strategies. These larvae are phenotypically varied across their range with predominantly black larvae (southeastern USA and California) and red larvae (western Texas, Arizona) occurring in different regions. Two years of field observations in south Texas indicate that the proportion of red larvae increases with increasing daily temperatures as the growing season progresses. Larvae were also observed to shift their microhabitats by climbing on non-host vegetation and avoided excessive heat in their feeding microhabitat. Larvae of ten half-sib families from populations in south Texas and California, reared under different temperature regimes in common garden experiments, exhibited plasticity in larval phenotype, with larvae from both populations producing the red phenotype at temperatures greater than 30°C and maintaining the black phenotype at cooler temperatures. However, larvae from Texas were more tolerant of higher temperatures, showing no decrease in growth rate in the highest temperature (maximum seasonal temperature) treatment, compared to the California population. In a field experiment, black larvae were found to have higher body temperatures when exposed to sunlight compared to red larvae. These results suggest that microhabitat shifts and the color polyphenism observed in pipevine swallowtail larvae may be the adaptive strategies that enable larvae to avoid critical thermal maximum temperatures.     

Geographical variation in wing polymorphism of the waterstrider Aquarius najas (Heteroptera,Gerridae)

The waterstrider Aquarius najas is wingless in Northern Europe, while winged individuals occur frequently in Central and Southern Europe. To test if the latitudinal difference is genetically controlled, we collected mature individuals from 10 different populations and raised their offspring in ‘common garden’ laboratory conditions. Half of these populations were from southern and the other half from central Finland. Daylength and temperature do influence wing development among other species of waterstriders, and thus we maintained a similar short daylength and warm conditions for all populations. These conditions should be favourable for wing development in general. Among laboratory-bred individuals several winged individuals appeared, and their proportion varied between populations. The relative frequency of winged individuals was highest in the southern populations. Thus, apart from phenotypic plasticity there seems to be some genetic control over the occurrence of wings, and the latitudinal trend coincides with the direction in natural populations over a larger European scale. Overwinter survival in our laboratory conditions was higher among the wingless individuals. The survival cost may explain why the proportion of winged individuals was lower in the northern populations with more extreme overwintering conditions than in the southern ones.     

Across-Shelf Transport of Bivalve Larvae: Can the Interface between a Coastal Current and Inshore Waters Act as an Ecological Barrier to Larval Dispersal?

Using an integrated physical and biological approach, we examined across-shelf advection and exchange and the associated transport of bivalve larvae in the presence of a strong coastal current separated from the coast by a stratified inshore environment. We tested the hypothesis that the interface of the coastal current and inshore waters can act as an ecological barrier to across-shelf transport of larvae but can be overcome by wind- or tidally-induced transport. Our study region in the Gulf of Maine encompasses a coastal current that diverges from the coast as it moves downshelf. The region inshore of this current is home to several species that exhibit limited recruitment in spite of extensive upshelf larval sources. Analysis of surface water temperatures and wind velocities revealed episodic decreases in temperature along the coast correlated with alongshelf (but not upwelling) winds, indicating wind-forced onshore movement of the cold coastal current. Such wind-driven onshore migrations are more common along the northern portion of the study region where the coastal current is near the coast, tidal currents are strong, and wind directions are more conducive to onshore migration, but rarer further south where the interface between inshore waters and the coastal current is further offshore and suitable wind events are less common. The distribution of bivalve larvae was consistent with the physical measurements. There was little across-shelf variation in larval abundance where the current abuts the coast, indicating strong across-shelf exchange of larvae, but strong across-shelf variation in larval density where the stratified inshore waters separate the current from the coast, indicating weak across-shelf transport of larvae. Our results suggest that the interface between the coastal current and inshore waters may constitute a major ecological barrier to larval dispersal in the southern part of the region that may only be overcome by rare, strong wind-forced events.     

Countergradient vs. cogradient variation in growth and diapause in a lichen‐feeding moth,Eilema depressum (Lepidoptera: Arctiidae)

Separating genetic and environmental causes of the latitudinal differences among populations is crucial when evaluating the potential for microevolutionary responses to the changing environment. We studied among‐population and environmental components of variation in several life‐history traits of a lichen‐feeding moth Eilema depressum when offspring of replicate Swiss and Finnish females were reared in a common‐garden factorial experiment. A partial second generation was produced only among Swiss larvae, more likely so at higher temperature regime and higher host quality, and more frequently among the offspring of particular females. Growth rates of larvae that chose the diapause development were higher in northern individuals. Our results thus reveal adaptive differences between latitudinal populations in studied life‐history traits, allowing to expect rapid adaptation of the species to further environmental changes. In contrast, invariable responses of the growth rates of the larvae to temperature and host quality support the idea that some basic parameters of insect growth show a high degree of evolutionary conservatism.     

Fecundity,autogeny, and the larval environment of the pitcher-plant mosquito,Wyeomyia smithii

Summary Pupae and fourth instar larvae of a southern (30°N, Alabama, USA) population of Wyeomyia smithii Coq. (Diptera: Culicidae) were collected from pitcher plants. Adults which emerged were maintained without food then dissected to determine their egg clutch size. Among females which matured eggs, fecundities were negatively correlated with larval densities in individual pitchers. The mean autogenous fecundity of the overwintering generation did not differ from a summer sample. Adults unable to mature eggs comprised 6–39% of samples, depending on whether collected as pupae or fourth instar larvae. Fecundity was negatively correlated with time to adult eclosion among larvae maintained on unrenewed pitcher contents in the laboratory.Cohorts from this population were reared in artificial containers from egg hatch to adulthood at a single density and a superior or inferior diet. On the superior larval diet, all females survived to reproductive age, and all but one (>99%) produced eggs autogenously. On the inferior diet, survivorship to adult eclosion was significantly less, a high proportion of females died before reaching reproductive age, and only 19% of survivors matured eggs without blood. Protracted larval development induced by the inferior diet did not influence the probability of autogeny among females that survived to reproductive maturity.The relationship between larval environment and reproductive strategies is contrasted across the geographic range of W. smithii. Bloodfeeding occurs among southern populations where density dependent constraints on preimaginal growth are constantly severe. The loss of hematophagy among northern populations may have been facilitated by periods of density independent larval growth.Institute of Food and Agricultural Sciences, University of Florida, Experimental Station Journal Series No. 3645     

Countergradient variation in behaviour,growth and lipid storage in Atlantic cod; environmental and genetic effects

We housed offspring from northern (70° N) and southern (60° N) coastal cod (Gadus morhua) together in a ‘common garden’ rearing experiment at a temperature and light regime representative of the southern population. Through a more active feeding behaviour and a higher success, the northern cod achieved a larger food share and a higher growth rate and condition than their southern conspecifics. This is contrary to what was demonstrated by field data of fish from their natural habitats. The northern cod also allocated more energy to the liver throughout the experiment. Our results agree with the theory of countergradient variation, suggesting that genetic influences on growth and condition have been opposed by environmental constraints in their natural habitat. The observation that the offspring from these populations differ in behavior and growth when housed together support the idea that the growth response to selection would be through a behavioral response. The field data suggest that density‐dependent population process and high juvenile density relative to prey limit the growth and condition in the wild and not necessarily the length of the growth season per se as assumed in the literature. The topographic distance (over 2000 km) limit mixing of early life stages of cod from the northern and southern population, and the different environmental stimuli (seasonality, temperature, food‐web interactions and habitat heterogeneity) in north and south are likely to evolve genetic differences.     

Latitudinal variation in resistance and tolerance to herbivory in the perennial herb Lythrum salicaria is related to intensity of herbivory and plant phenology

Both the length of the growing season and the intensity of herbivory often vary along climatic gradients, which may result in divergent selection on plant phenology, and on resistance and tolerance to herbivory. In Sweden, the length of the growing season and the number of insect herbivore species feeding on the perennial herb Lythrum salicaria decrease from south to north. Previous common‐garden experiments have shown that northern L. salicaria populations develop aboveground shoots earlier in the summer and finish growth before southern populations do. We tested the hypotheses that resistance and tolerance to damage vary with latitude in L. salicaria and are positively related to the intensity of herbivory in natural populations. We quantified resistance and tolerance of populations sampled along a latitudinal gradient by scoring damage from natural herbivores and fitness in a common‐garden experiment in the field and by documenting oviposition and feeding preference by specialist leaf beetles in a glasshouse experiment. Plant resistance decreased with latitude of origin, whereas plant tolerance increased. Oviposition and feeding preference in the glasshouse and leaf damage in the common‐garden experiment were negatively related to damage in the source populations. The latitudinal variation in resistance was thus consistent with reduced selection from herbivores towards the northern range margin of L. salicaria. Variation in tolerance may be related to differences in the timing of damage in relation to the seasonal pattern of plant growth, as northern genotypes have developed further than southern have when herbivores emerge in early summer.