Life‐history traits related to diapause in univoltine and bivoltine populations of Ypthima multistriata (Lepidoptera: Satyridae) inhabiting similar latitudes

In most temperate insects, diapause strategies and voltinism generally exhibit latitudinal clines, supporting the concept that they represent adaptations to climate. In contrast, in the satyrine butterfly Ypthima multistriata Butler, local populations with different voltinism patterns are geographically intermingled, suggesting that life‐history traits related to diapause may differ even between geographically and phylogenetically close populations. In this study, we experimentally examined the critical photoperiod for diapause induction and the larval developmental period in two univoltine and two bivoltine populations of Y. multistriata, all of which inhabit virtually the same latitude (34.652–34.750°N). We found that the critical photoperiod for diapause induction was longer in the univoltine populations than in the bivoltine populations. Moreover, the larval period under the long day length treatment was different among populations in both sexes, although significant differences were also detected between populations with the same voltinism. These results indicate that in Y. multistriata, life‐history traits related to diapause can not be attributed merely to climatic conditions such as temperature or day length, which depend largely on latitude. Therefore, we suggest that biotic elements, such as leaf toughness, as well as abiotic elements should be taken into account in attempts to explain the enigmatic pattern of geographic variation in the diapause strategies of Y. multistriata.     

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.     

Time constraint effects on phenology and life history synchrony in a damselfly along a latitudinal gradient

In organisms with complex life cycles living in seasonal environments, the synchronisation of phenological events is important from the ecological and evolutionary perspectives. Life history transitions should be synchronised to a greater degree at northern latitudes. We quantified hatching and emergence timing and synchrony in the obligate univoltine damselfly Lestes sponsa along a latitudinal gradient covering its entire north–south range in Europe. In our first experiment, populations from different latitudes were grown in separate climate chambers simulating temperature and photoperiod conditions occurring at their sites of origin. Northern populations expressed early and high synchronous hatching and emergence, central populations intermediate, and southern populations late and low synchronous hatching and emergence. This pattern was expressed at both population and full‐sibling family levels, indicating stronger selection for timing and synchronisation in the north compared to the south. In our second experiment, populations from all latitudes were reared in conditions simulating an average temperature and photoperiod over the latitudinal gradient. Interestingly, the pattern of timing and synchronisation was reversed with respect to latitude when compared to the pattern shown in the first experiment, indicating the importance of environmental factors in shaping phenological events. Our results indicate strong selection for timing and synchronisation of life history events at northern latitudes, caused by time constraints. Our results also show that it is important to use as natural conditions as possible in experiments on life history shifts in organisms with complex life cycles in order to achieve a correct understanding of these shifts.     

Females show greater changes in wing colour with latitude than males in the green‐veined white butterfly,Pieris napi (Lepidoptera: Pieridae)

In butterflies, wing colour may simultaneously be under sexual selection in the context of mating selection and natural selection in the context of thermoregulation. In the present study, we collected mated females of the green‐veined white butterfly (Pieris napi) from locations spanning 960 km of latitude across Fennoscandia, and investigated sex‐specific latitudinal wing colour variation in their offspring raised under identical conditions. We measured wing colour characteristics, including reflectance at wavelengths 300–700 nm and the degree of wing melanization. At all latitudes, females reflected more light in the short wavelengths (< 400 nm) and less in the long wavelengths (> 450 nm), and they were more melanized than males. However, female wing colour varied more with latitude than that of males. Among females, long wavelength reflectance decreased, whereas short wavelength reflectance and melanization increased, towards the north. By contrast, among males, latitudinal variation was found only in the ventral hindwing melanization. These results are consistent with the idea that the balance between natural and sexual selection acting on wing colour changes with latitude differently in males than females. The dark wing colour of females in the north may be a thermoregulatory adaptation, although males may be constrained from evolving the dark dorsal wing colour favoured by natural selection because of constant sexual selection across latitudes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Opposing clines for high and low temperature resistance in Drosophila melanogaster

In insects, species comparisons suggest a weak association between upper thermal limits and latitude in contrast to a stronger association for lower limits. To compare this to latitudinal patterns of thermal responses within species, we considered latitudinal variation in heat and cold resistance in Drosophila melanogaster. We found opposing clines in resistance to these temperature extremes in comparisons of 17–24 populations from coastal eastern Australia. Knockdown time following heat shock increased towards the tropics, whereas recovery time following cold shock decreased towards temperate latitudes. Mortality following cold shock also showed a clinal pattern. Clinal associations with latitude were linear and related to minimum temperatures in the coldest month (for cold resistance) and maximum temperatures in the warmest month (for heat resistance). This suggests that within species both high and low temperature responses can vary with latitude as a consequence of direct or indirect effects of selection.     

The roles of pre‐ and post‐hatching growth rates in generating a latitudinal cline of body size in the eastern fence lizard (Sceloporus undulatus)

Countergradient variation in norms of reaction can dampen the direct effects of environmental influences on phenotypic traits, allowing phenotypic similarity among populations despite exposure to different environmental conditions. Such norms of reaction may occur at any phase of the life‐history (e.g. growth rates during both embryonic and postembryonic stages may influence geographical variation in adult body size). We collected gravid female lizards (Sceloporus undulatus) from northern (Indiana), central (Mississippi), and southern (Florida) populations, spanning almost the full latitudinal range of the species. Adult females from the southern population were smaller. Intrinsic growth rates of hatchlings were higher for the central population than for the other two populations. This pattern does not parallel the countergradient variation previously found in embryonic developmental rates among these populations. Earlier hatching enhanced survival rates of juveniles to a similar degree among populations, although juvenile survival rates in the field generally increase with latitude in this species. Our data reveal geographical variation in the ways in which intrinsic developmental/growth rates and survival shift during ontogeny, and suggest that latitudinal patterns in adult body size (such as Bergmann's rule) can result from both faster growth, and longer periods of growth. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 202–209.     

Genetic and phenotypic sources of life history variation along a cline in voltinism in the cricket Allonemobius socius

Clinal variation in life histories can be genetically based, resulting from selection imposed by different environments, or it may be due to the differential expression of phenotypically plastic traits. We examined the cline in voltinism in the egg-diapausing cricket Allonemobius socius, with populations spanning the switch from a univoltine to a bivoltine phenology. A common garden experiment was employed, using environments that mimicked photoperiod and temperature conditions found in the field. There were only small differences in development time among populations, and the difference in phenology observed in the field is likely due to clinal variation in the length of the growing season. We found large genetically-based differences in the reaction norm for egg diapause that were further magnified by environmental cues. The synergism of genetic and environmental effects was an example of cogradient selection. In the zone of transition between phenologies, voltinism appeared to be a conditional strategy, rather than a genetic polymorphism. First-generation females from this area can lay either direct-developing or diapause eggs depending on the likelihood that a second generation will have sufficient time to develop. For this species, the cline in voltinism is the result of a combination of environmental effects on development, and genetic and environmental influences on egg diapause propensity.     

Latitudinal and voltinism compensation shape thermal reaction norms for growth rate

Latitudinal variation in thermal reaction norms of key fitness traits may inform about the response of populations to climate warming, yet their adaptive nature and evolutionary potential are poorly known. We assessed the contribution of quantitative genetic, neutral genetic and environmental effects to thermal reaction norms of growth rate for populations of the damselfly Ischnura elegans. Among populations, reaction norms differed primarily in elevation, suggesting that time constraints associated with shorter growth seasons in univoltine, high-latitude as well as multivoltine, low-latitude populations selected for faster growth rates. Phenotypic divergence among populations is consistent with selection rather than drift as Q(ST) was greater than F(ST) in all cases. Q(ST) estimates increased with experimental temperature and were influenced by genotype by environment interactions. Substantial additive genetic variation for growth rate in all populations suggests that evolution of trait means in different environments is not constrained. Heritability of growth rates was higher at high temperature, driven by increased genetic rather than environmental variance. While environment-specific nonadditive effects also may contribute to heritability differences among temperatures, maternal effects did not play a significant role (where these could be accounted for). Genotype by environment interactions strongly influenced the adaptive potential of populations, and our results suggest the potential for microevolution of thermal reaction norms in each of the studied populations. In summary, the observed latitudinal pattern in growth rates is adaptive and results from a combination of latitudinal and voltinism compensation. Combined with the evolutionary potential of thermal reaction norms, this may affect populations' ability to respond to future climate warming.     

A test of the latitudinal defense hypothesis: herbivory, tannins and total phenolics in four North American tree species

It is widely believed that insect herbivory is less intense at higher latitudes, due to winter mortality which would tend to keep insect herbivores from reaching density-limitation of their populations. One prediction of this theory is that plants should tend to be better defended at lower latitudes. Here we investigated latitudinal trends in herbivory and tannins, in four species of common North American trees. Our comparisons spanned 15° of latitude in Acer rubrum, Fagus grandifolia, and Quercus alba, and 10° latitude in Liquidambar styraciflua. Sun leaves on forest edges were sampled, at phenologically equivalent times of year. Analysis revealed significant differences between populations, including those at similar latitudes, but no significant latitudinal trend in herbivory, condensed and hydrolyzable tannins, or total phenolics measured as Folin–Denis reactives in any of the four species. Our findings contradict the theory that low latitude plants are better defended, in that lower latitude populations of the four tree species showed no greater amounts of phenolics. The possible implications for community ecology are discussed. Jonathan M. Adams, Brian Rehill, and Yangjian Zhang are equal joint lead authorship on this paper.     

Are European White Butterflies Aposematic?

It has been suggested that the white coloration of Pieridae butterflies is a warning signal and therefore all white Pieridae could profit from a mimetic resemblance. We tested whether green-veined white (Pieris napi) and orange-tip (Anthocharis cardamines) butterflies benefit from white coloration. We compared their relative acceptability to wild, adult pied flycatchers (Ficedula hypoleuca) by offering live A. cardamines and P. napi together with two non-aposematic butterflies on the tray attached to birds' nesting boxes. Experienced predators equally attacked white and non-white butterflies, and the order of attack among the Pieridae was random. If anything, there was a slight indication that the female A. cardamines was the least favoured prey. Since birds did not avoid white coloration, we compared the palatability of these two species against known palatable and unpalatable butterflies by presenting them to great tits (Parus major). Pieris brassicae, which has been earlier described as unpalatable, was also included in the palatability test. However, there were no significant differences in the palatability of the butterflies to birds, and even P. brassicae was apparently palatable to the great tits. Our results do not unambiguously support the hypothesis that the white coloration of Pieridae would signal unpalatability. Nevertheless, in our last experiment, pied flycatchers often rejected or left untouched free flying P. napi and A. cardamines. This suggests that other features in a more natural situation, such as the agile flight pattern or odours might still make them unprofitable to birds. This revised version was published online in July 2006 with corrections to the Cover Date.     

Differential effects of short term winter thermal stress on diapausing tiger swallowtail butterflies (Papilio spp.)

Abstract It is generally thought that insects inhabiting lower latitudes are more severely impacted by changes in their thermal environment than are high latitude species. This is attributed to the wider range of temperatures to which high‐latitude species are exposed. By contrast, low‐latitude species have typically evolved in more thermally stable environments with a narrower range of temperature variation. However, deviation from this pattern can occur and here we report that under variable winter conditions a higher latitude species may be more sensitive to thermal variation than its lower latitude sister species. Using split broods, we examined the survival and adult emergence success of diapausing pupae of Papilio canadensis and P. glaucus, as well as a unique, recombinant hybrid population (“late‐flight”) to short periods of mid‐winter cold and heat stress. Our results indicate that the higher latitude, univoltine populations (P. canadensis and late‐flights) exhibit lower pupal survival than the lower latitude, facultative diapauser (P. glaucus) for all mid‐winter thermal stress treatments, both high and low. Size differences alone do not appear to account for the observed differences in survival or metabolic costs in these three phenotypes, as late‐flight individuals are similar in size to P. glaucus. We attribute the observed differences in survival and weight loss to potential metabolic differences and variation in the intensity of diapause, in addition to divergent adaptation to winter precipitation levels (e.g. snow cover) and the influences this may have on microhabitat temperature moderation.     

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.     

Variation in plant thermal reaction norms along a latitudinal gradient – more than adaptation to season length

Little is known about the extent to which observed phenological responses to changes in climate are the result of phenotypic plasticity or genetic changes. We also know little about how plasticity, in terms of thermal reaction norms, vary spatially. We investigated if the thermal reaction norms for flower development of five crucifer species (Brassicaceae) differed among three regions along a south–north latitudinal gradient in replicated experiments. The mean response (elevation) of thermal reaction norms of flowering differed among regions in all study species, while sensitivity of flower development to temperature (slope) differed in only one of the species. Differences in mean responses corresponded to cogradient patterns in some species, but countergradient patterns in other. This suggests that differences among regions were not solely the result of adaptation to differences in the length of the growing season, but that other factors, such as herbivory, play an important role. Differences in development rate within species were mainly explained by variation in early phases of bud formation in some species but by variation in later phases of bud formation in other species. The differences in latitudinal patterns of thermal reaction norms among species observed in this study are important, both to identify agents of selection and to predict short‐ and long‐term responses to a warming climate.     

Latitudinal patterns in the phenological responses of leaf colouring and leaf fall to climate change in Japan

Aim  To estimate the potential effect of global climate change on the phenological responses of plants it is necessary to estimate spatial variations at larger scales. However, previous studies have not estimated latitudinal patterns in the phenological response directly. We hypothesized that the phenological response of plants varies with latitude, and estimated the phenological response to long-term climate change using autumn phenological events that have been delayed by recent climate change.
Location  Japan.
Methods  We used a 53-year data set to document the latitudinal patterns in the climate responses of the timing of autumn leaf colouring and fall for two tree species over a wide range of latitudes in Japan (31 to 44° N). We calculated single regression slopes for leaf phenological timing and air temperature across Japan and tested their latitudinal patterns using regression models. The effects of latitude, time and their interaction on the responses of the phenological timings were also estimated using generalized linear mixed models.
Results  Our results showed that single regression slopes of leaf phenological timing and air temperature in autumn were positive at most stations. Higher temperatures can delay the timing of leaf phenology. Negative relationships were found between the phenological response of leaves to temperature and latitude. Single regression slopes of the phenological responses at lower latitudes were larger than those at higher latitudes.
Main conclusions  We found negative relationships between leaf phenological responsiveness and latitude. These findings will be important for predicting phenological timing with global climate change.     

Phenological matching rather than genetic variation in host preference underlies geographical variation in host plants used by orange tip butterflies

An insect species that shows variation in host species association across its geographical range may do so either because of local adaptation in host plant preference of the insect or through environmentally or genetically induced differences in the plants, causing variation in host plant suitability between regions. In the present study, we experimentally investigate the host plant preference of Anthocharis cardamines (orange tip butterfly) in two populations from the UK and two from Sweden. Previous reports indicate that A. cardamines larvae are found on different host plant species in different regions of the UK, and some variation has been reported in Sweden. Host plant choice trials showed that females prefer to oviposit on plants in an earlier phenological stage, as well as on larger plants. When controlling for plant phenological stage and size, the host species had no statistically significant effect on the choice of the females. Moreover, there were no differences in host plant species preference among the four butterfly populations. Based on our experiment, the oviposition choice by A. cardamines mainly depends on the phenological stage and the size of the host plant. This finding supports the idea that the geographical patterns of host–plant association of A. cardamines in the UK and Sweden are consequences of the phenology and availability of the local hosts, rather than regional genetic differences in the host species preference of the butterfly.     

Explaining the sawtooth: latitudinal periodicity in a circadian gene correlates with shifts in generation number

Many temperate insects take advantage of longer growing seasons at lower latitudes by increasing their generation number or voltinism. In some insects, development time abruptly decreases when additional generations are fit into the season. Consequently, latitudinal ‘sawtooth’ clines associated with shifts in voltinism are seen for phenotypes correlated with development time, like body size. However, latitudinal variation in voltinism has not been linked to genetic variation at specific loci. Here, we show a pattern in allele frequency among voltinism ecotypes of the European corn borer moth (Ostrinia nubilalis) that is reminiscent of a sawtooth cline. We characterized 145 autosomal and sex‐linked SNPs and found that period, a circadian gene that is genetically linked to a major QTL determining variation in post‐diapause development time, shows cyclical variation between voltinism ecotypes. Allele frequencies at an unlinked circadian clock gene cryptochrome1 were correlated with period. These results suggest that selection on development time to ‘fit’ complete life cycles into a latitudinally varying growing season produces oscillations in alleles associated with voltinism, primarily through changes at loci underlying the duration of transitions between diapause and other life history phases. Correlations among clock loci suggest possible coupling between the circadian clock and the circannual rhythms for synchronizing seasonal life history. We anticipate that latitudinal oscillations in allele frequency will represent signatures of adaptation to seasonal environments in other insects and may be critical to understanding the ecological and evolutionary consequences of variable environments, including response to global climate change.     

Latitudinal variation in plant chemical defences drives latitudinal patterns of leaf herbivory

A long‐standing paradigm in ecology holds that herbivore pressure and thus plant defences increase towards lower latitudes. However, recent work has challenged this prediction where studies have found no relationship or opposite trends where herbivory or plant defences increase at higher latitudes. Here we tested for latitudinal variation in herbivory, chemical defences (phenolic compounds), and nutritional traits (phosphorus and nitrogen) in leaves of a long‐lived tree species, the English oak Quercus robur. We further investigated the underlying climatic and soil factors associated with such variation. Across 38 populations of Q. robur distributed along an 18° latitudinal gradient, covering almost the entire latitudinal and climatic range of this species, we observed strong but divergent latitudinal gradients in leaf herbivory and leaf chemical defences and nutrients. As expected, there was a negative relationship between latitude and leaf herbivory where oak populations from lower latitudes exhibited higher levels of leaf herbivory. However, counter to predictions there was a positive relationship between leaf chemical defences and latitude where populations at higher latitudes were better defended. Similarly, leaf phosphorus and nitrogen increased with latitude. Path analysis indicated a significant (negative) effect of plant chemical defences (condensed tannins) on leaf herbivory, suggesting that the latitudinal gradient in leaf herbivory was driven by an inverse gradient in defensive investment. Leaf nutrients had no independent influence on herbivory. Further, we found significant indirect effects of precipitation and soil porosity on leaf herbivory, which were mediated by plant chemical defences. These findings suggest that abiotic factors shape latitudinal variation in plant defences and that these defences in turn underlie latitudinal variation in leaf herbivory. Overall, this study contributes to a better understanding of latitudinal variation in plant–herbivore interactions by determining the identity and modus operandi of abiotic factors concurrently shaping plant defences and herbivory.     

Evolution,plasticity and evolving plasticity of phenology in the tree species Alnus glutinosa

Both traits and the plasticity of these traits are subject to evolutionary change and therefore affect the long‐term persistence of populations and their role in local communities. We subjected clones from 12 different populations of Alnus glutinosa, located along a latitudinal gradient, to two different temperature treatments, to disentangle the distribution of genetic variation in timing of bud burst and bud burst plasticity within and among genotypes, populations, and regions. We calculated heritability and evolvability estimates for bud burst and bud burst plasticity and assessed the influence of divergent selection relative to neutral drift. We observed higher levels of heritability and evolvability for bud burst than for its plasticity, whereas the total phenological heritability and evolvability (i.e. combining timing of bud burst and bud burst plasticity) suggest substantial evolutionary potential with respect to phenology. Earlier bud burst was observed for the low‐latitudinal populations than for the populations from higher latitudes, whereas the high‐latitudinal populations did not show the expected delayed bud burst. This countergradient variation can be due to evolution towards increased phenological plasticity at higher latitudes. However, because we found little evidence for adaptive differences in phenological plasticity across the latitudinal gradient, we suggest differential frost tolerance as the most likely explanation for the observed phenological patterns in A. glutinosa.     

Phylogeography reveals latitudinal population structure in the common herb Plantago coronopus

The increase in gene diversity from high to low latitudes is a widely recognized biogeographical pattern, often shaped by differential effects of Late Quaternary climatic changes. Here, we evaluate the effects of Pleistocene climatic changes from northern Europe to North Africa and their implications on the population differentiation of the widespread, short‐lived herb Plantago coronopus. We used amplified fragment length polymorphism to investigate the population structure and phylogeography of P. coronopus in 273 individuals from 29 populations covering its complete latitudinal range. Although Bayesian clustering, principal coordinates analysis and a consensus UPGMA tree were not fully congruent, two well‐supported clades, associated with distinct latitudinal zones (northern Europe and the Mediterranean region), were revealed as a general pattern. Moreover, populations from the western Atlantic edge and, to a lesser extent, the central Mediterranean region exhibited signs of admixture, suggesting secondary contacts. The admixed populations in the western Atlantic and central Mediterranean are geographically intermediate between the northern and southern lineages. The northernmost lineage exhibited low genetic diversity, a clear sign of a recent colonization. In contrast, populations from the southernmost part of the range showed the highest level of genetic diversity, indicating possible refugia for the species during the Quaternary ice ages. Overall, our study allows spatial structure of the genetic variation of a widespread herb across its latitudinal range to be disentangled and provides insights into how past climatic history influences present genetic patterns. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 618–634.     

Latitudinal variation in genetic divergence of populations and the potential for future speciation

The increase in biological diversity with decreasing latitude is widely appreciated but the cause of the pattern is unknown. This pattern reflects latitudinal variation in both the origin of new species (cladogenesis) and the number of species that coexist. Here we address latitudinal variation in species origination, by examining population genetic processes that influence speciation. Previous data suggest a greater number of speciation events at lower latitudes. If speciation events occur more frequently at lower latitudes, we predicted that genetic divergence among populations within species, an important component of cladogenesis, should be greater among lower latitude populations. We tested this prediction using within-species patterns of mtDNA variation across 60 vertebrate species that collectively spanned six continents, two oceans, and 119 degrees latitude. We found greater genetic divergence of populations, controlling for geographic distance, at lower latitudes within species. This pattern remained statistically significant after removing populations that occur in localities previously covered by continental glaciers during the last glaciation. Results suggest that lower latitude populations within species exhibit greater evolutionary independence, increasing the likelihood that mutation, recombination, selection, and/or drift will lead to divergence of traits important for reproductive isolation and speciation. Results are consistent with a greater influence of seasonality, reduced energy, and/or glacial (Milankovitch) cycles acting on higher latitude populations, and represent one of the few tests of predictions of latitudinal variation in speciation rates using population genetic data.