Clinal variation in fitness related traits in tropical drosophilids of the Indian subcontinent

In the last two decades a considerable number of studies have described geographic variation in morphometric, physiological and behavioral traits of over a dozen species of wild south-Asian drosophild fruit flies. Due to strong latitudinal and altitudinal gradients this region is highly diverse with respect to climatic conditions, making it important for studies in evolutionary ecology. In this review, we examine spatial heterogeneity across the Indian subcontinent in almost all of the traits previously investigated (body weight, desiccation tolerance, pigmentation, copulation duration, fecundity, ovariole number, wing length, alcohol dehydrogenase fast allele frequency, rate of water loss and starvation resistance). We find a linear correlation between trait variation and latitude. Our data suggest that a single climatic component, the coefficient of variance of monthly temperature averages, which is strongly correlated with latitude, explains a large proportion of variation in the traits investigated.     

Nonrandom variation of morphological traits across environmental gradients in a land snail

Morphological variation is often attributed to differential adaptations to diverse habitats, but adaptations to a similar environment do not necessarily result in similar phenotypes. Adaptations for water and heat budget are crucial for organisms living in arid habitats, and in snails, variation in shell morphology has been frequently attributed to selection by stressful environmental factors. However, their phenotypic divergence often is not accompanied by a relevant niche differentiation and consistent relationships with environmental correlates are lacking. In the pulmonate genus Albinaria, there is great size and shape variation between and within species, and there are two major shell sculpture morphotypes, ribbed and smooth. We used 62 populations of 28 Albinaria species, taking into account their phylogeny, to examine the variation of shell traits (sculpture, size, shape), their effect on water and heat budget, and their association with geographical and climatic gradients. We found unambiguous size and shape discrimination between the two morphotypes. Ribbed shells are lighter, taller, and slimmer and have a smaller aperture than the smooth ones. Moreover, significant correlations between shell traits and heat/moisture budget and climate/geography were revealed. Ribbed and taller shells retain more water on their shell surface, and on the other hand, smooth shells exhibit lower water permeability. Therefore, two strategies are being used to prevent water loss, active retention or resistance to loss. Consequently, different alternative solutions evolved and were retained as responses to the same stressful factor by the two distinct shell morphotypes. Larger shells occur in southern latitudes, mostly on islands, and at sites where there is a shortage of rainfall. Therefore, the variation of the examined traits is nonrandom with respect to location and to climate and their evolution can be attributed to selection by environmental factors, with water availability being the key driving agent of body-size variation.     

Novel variation associated with species range expansion

When species shift their ranges to track climate change, they are almost certain to experience novel environments to which they are poorly adapted. Otaki and co-workers document an explosion of wing pattern variation accompanying range expansion in the pale grass blue butterfly. This pattern can be replicated in the laboratory using artificial selection on cold shocked pupae, at temperature extremes typical of recently colonized environments. We discuss how this phenotypic plasticity may be associated with successful colonization and how significant local adaptation is likely to re-establish developmental control. Integrating knowledge of trait plasticity into current genetic models of adaptation is central to our understanding of when and where a colonising population will be able to persist and adapt in novel surroundings.     

Factors influencing diet variation in a generalist predator across its range distribution

Factors linked with intraspecific variation in trophic diversity are still poorly understood in generalist species like the Montagu’s harrier (Circus pygargus) but may have important implications for conservation management at a wide scale. We described geographic patterns of Montagu’s harrier diet across Eurasia, gathering diet data from 30 studies in 41 areas from 11 countries. We grouped prey as invertebrates, reptiles, small mammals, large mammals, eggs, small birds and large birds, and calculated the contribution of each prey type to the diet (as % biomass) and Shannon’s Diversity Index for each study site. We analysed qualitative estimates of prey abundance in relation to latitude and longitude, then diet composition in relation to habitat of the study area and prey abundance estimates. Diet diversity of Montagu’s harriers increased from north to south, while abundance of all prey groups other than small mammals showed the opposite trends. Agricultural areas in northern latitudes seemed to hold high densities of small mammals, but low densities of alternative prey. Overall, birds were the main prey in most of Montagu’s harrier’s distribution range, although the relative importance of each prey type in the diet was significantly explained by its local abundance and habitat, confirming the opportunistic foraging strategy of this raptor species. Consumption of mammals was an exception to this trend, being negatively associated with the abundance of alternative prey, suggesting that this prey is not preferred. Trophic diversity in this species could be influenced by land-use changes through variations in the abundance and availability of prey, which could impact its population dynamics. This may be particularly important for northern populations of Montagu’s harriers breeding in agricultural habitats, where trophic diversity is already low.     

Life-history traits of lake plankton species may govern their phenological response to climate warming

A prominent response of temperate aquatic ecosystems to climate warming is changes in phenology – advancements or delays in annually reoccurring events in an organism's life cycle. The exact seasonal timing of warming, in conjunction with species-specific life-history events such as emergence from resting stages, timing of spawning, generation times, or stage-specific prey requirements, may determine the nature of a species' response. We demonstrate that recent climate-induced shifts in the phenology of lake phytoplankton and zooplankton species in a temperate eutrophic lake (Müggelsee, Germany) differed according to differences in their characteristic life cycles. Fast-growing plankton in spring (diatoms, Daphnia ) showed significant and synchronous forward movements by about 1 month, induced by concurrent earlier ice break-up dates (diatoms) and higher spring water temperature ( Daphnia ). No such synchrony was observed for slow-growing summer zooplankton species with longer and more complex life cycles (copepods, larvae of the mussel Dreissena polymorpha ). Although coexisting, the summer plankton responded species specifically to seasonal warming trends, depending on whether the timing of warming matched their individual thermal requirements at decisive developmental stages such as emergence from diapause (copepods), or spawning ( Dreissena ). Others did not change their phenology significantly, but nevertheless, increased in abundances. We show that the detailed seasonal pattern of warming influences the response of phyto- and zooplankton species to climate change, and point to the diverse nature of responses for species exhibiting complex life-history traits.     

Natural variation for drought-response traits in the Mimulus guttatus species complex

Soil moisture is a key factor affecting plant abundance and distribution, both across and within species. In response to water limitation, plants have evolved numerous morphological, physiological, and phenological adaptations. In both well-watered and water-limited conditions, we identified considerable natural variation in drought-related whole-plant and leaf-level traits among closely related members of the Mimulus guttatus species complex that occupy a diversity of habitats in the field. The self-fertilizing Mimulus nasutus and serpentine-endemic Mimulus nudatus demonstrated the overall greatest tolerance to soil water limitation, exhibiting the smallest reduction in seed set relative to well-watered conditions. This may be due in part to early flowering, faster fruit development, and low stomatal density. In contrast, flowering of coastal M. guttatus was so delayed that it precluded any seed production in water-limited conditions. This range of phenotypic responses to soil water deficit in Mimulus, coupled with developing genomic resources, holds considerable promise for identifying genomic variation responsible for adaptive responses to soil water availability.     

Phenological response of sea turtles to environmental variation across a species' northern range

Variations in environmental parameters (e.g. temperature) that form part of global climate change have been associated with shifts in the timing of seasonal events for a broad range of organisms. Most studies evaluating such phenological shifts of individual taxa have focused on a limited number of locations, making it difficult to assess how such shifts vary regionally across a species range. Here, by using 1445 records of the date of first nesting for loggerhead sea turtles (Caretta caretta) at different breeding sites, on different continents and in different years across a broad latitudinal range (25–39° ′N), we demonstrate that the gradient of the relationship between temperature and the date of first breeding is steeper at higher latitudes, i.e. the phenological responses to temperature appear strongest at the poleward range limit. These findings support the hypothesis that biological changes in response to climate change will be most acute at the poleward range limits and are in accordance with the predictions of MacArthur''s hypothesis that poleward range limit for species range is environmentally limited. Our findings imply that the poleward populations of loggerheads are more sensitive to climate variations and thus they might display the impacts of climate change sooner and more prominently.     

Clinal variation in Drosophila serrata for stress resistance and body size

Clines for size and stress resistance traits have been described for several Drosophila species and replicable clines across different species may indicate climatic selection. Here we consider clines in stress resistance traits in an Australian endemic species, D. serrata, by comparing levels of variation within and among isofemale lines initiated with flies collected from the eastern coast of Australia. We also consider clinical variation in chill coma recovery, a trait that has recently been shown to exhibit high levels of variation among Drosophila species. Patterns were compared with those in the cosmopolitan species D. melanogaster from the same area. Both desiccation and starvation resistance showed no clinical pattern despite heritable variation among isofemale lines. In contrast chill coma resistance exhibited a linear cline in the anticipated direction, resistance increasing with latitude. Body size was measured as wing length and body weight. Both traits showed geographic variation and strong non-linear clines with a sharp reduction in size in the tropics. These results are discussed in the context of climatic selection and evolutionary processes limiting species borders.     

Leaf traits determine the growth-survival trade-off across rain forest tree species

A dominant hypothesis explaining tree species coexistence in tropical forest is that trade-offs in characters allow species to adapt to different light environments, but tests for this hypothesis are scarce. This study is the first that uses a theoretical plant growth model to link leaf trade-offs to whole-plant performances and to differential performances across species in different light environments. Using data of 50 sympatric tree species from a Bolivian rain forest, we observed that specific leaf area and photosynthetic capacity codetermined interspecific height growth variation in a forest gap; that leaf survival rate determined the variation in plant survival rate under a closed canopy; that predicted height growth and plant survival rate matched field observations; and that fast-growing species had low survival rates for both field and predicted values. These results show how leaf trade-offs influence differential tree performance and tree species' coexistence in a heterogeneous light environment.     

Clinal variation in body and cell size in a widely distributed vertebrate ectotherm

Bergmanns rule states that, among conspecific populations, individuals are larger in cooler than in warmer environments as a consequence of selection related to heat conservation. Many of the most comprehensive assessments of Bergmanns rule to date have examined clinal patterns in body size among species assemblages. Our study is a more direct test of Bergmanns rule because we examine the pattern within a single, widely distributed species. We examined geographic variation in body and cell size in the spotted turtle (Clemmys guttata). Our analysis of 818 turtles collected from the entire range (45–28°N), indicated that body size increased with latitude; however, the relationship was driven by a population of large turtles at the northern extreme of the species range. When the northern population was removed from the analyses, Bergmanns rule was not supported, and the smallest turtles occurred near the central part of the species distribution. Recent literature has suggested that latitudinal clines in body size may simply be a physiological byproduct of the effects of temperature on cell division, resulting in larger cells, and hence larger organisms, from cooler temperatures. Measurements of the diameter of skin cells did not support the hypothesis that cell size increases with latitude and decreases with temperature in the spotted turtle, nor was there a significant relationship between body size and cell size. Our study suggests that neither Bergmanns rule nor cell size variation sufficiently explain the body size cline observed in the spotted turtle. We hypothesize that patterns in body size are related to variation in female size at maturity and reproductive cycles.     

Genetic variation and covariation of aphid life-history traits across unrelated host plants

A central paradigm of life-history theory is the existence of resource mediated trade-offs among different traits that contribute to fitness, yet observations inconsistent with this tenet are not uncommon. We previously found a clonal population of the aphid Myzus persicae to exhibit positive genetic correlations among major components of fitness, resulting in strong heritable fitness differences on a common host. This raises the question of how this genetic variation is maintained. One hypothesis states that variation for resource acquisition on different hosts may override variation for allocation, predicting strong fitness differences within hosts as a rule, but changes in fitness hierarchies across hosts due to trade-offs. Therefore, we carried out a life-table experiment with 17 clones of M. persicae, reared on three unrelated host plants: radish, common lambsquarters and black nightshade. We estimated the broad-sense heritabilities of six life-history traits on each host, the genetic correlations among traits within hosts, and the genetic correlations among traits on different hosts (cross-environment genetic correlations). The three plants represented radically different environments with strong effects on performance of M. persicae, yet we detected little evidence for trade-offs. Fitness components were positively correlated within hosts but also between the two more benign hosts (radish and lambsquarters), as well as between those and another host tested earlier. The comparison with the most stressful host, nightshade, was hampered by low survival. Survival on nightshade also exhibited genetic variation but was unrelated to fitness on other hosts. Acknowledging that the number of environments was necessarily limited in a quantitative genetic experiment, we suggest that the rather consistent fitness hierarchies across very different plants provided little evidence to support the idea that the clonal variation for life-history traits and their covariance structure are maintained by strong genotypexenvironment interactions with respect to hosts. Alternative explanations are discussed.     

The role of short-tongued insects in floral variation across the range of a style-dimorphic plant

Background and Aims

Heterostyly and related style polymorphisms are suitable model systems to evaluate the importance of functional pollinators in the maintenance of population variability. In Narcissus papyraceus different functional pollinators, incompatibility system and flower morphology have been proposed to influence the maintenance of polymorphism through their effect on disassortative mating. Here a test is done to find out if the visitation rate of long- versus short-tongued pollinators correlates with the morph ratio and if the latter is related to other flower traits of the species across its main geographic range.

Methods

Floral traits from 34 populations in the south-west of the Iberian Peninsula and in north-west Africa were measured, perianth variation was described and a comparison was made of allometric relationships between sex organs and floral tube. Correlations between pollinator guilds, stigma–anther separation of reciprocal morphs (our proxy for disassortative mating) and morph-ratio variation were analysed. Finally, the incompatibility system of the species in the northern and southern borders of its distribution are described.

Key Results

Flowers from southern populations were significantly larger than flowers from centre and northern populations. The abundance of short-styled plants decreased gradually with increasing distance from the core region (the Strait of Gibraltar), with these disappearing only in the northern range. Although there was a significant difference in stigma–anther separation among populations, morph ratio was not associated with reciprocity or floral tube length. Long-style morph frequency increased with short-tongued pollinator visitation rate. Populations from both edges of the distribution range were self-incompatible and within- and between-morph compatible.

Conclusions

The style morph ratio changed gradually, whereas perianth trait variation showed abrupt changes with two morphotypes across the range. The positive relationship between the visitation rate of short-tongued pollinators and the decrease of the short-style morph supports our initial hypothesis. The results highlight the importance of different pollinators in determining the presence of style polymorphism.     

Clinal variation for amino acid polymorphisms at the Pgm locus in Drosophila melanogaster

Clinal variation is common for enzymes in the glycolytic pathway for Drosophila melanogaster and is generally accepted as an adaptive response to different climates. Although the enzyme phosphoglucomutase (PGM) possesses several allozyme polymorphisms, it is unique in that it had been reported to show no clinal variation. Our recent DNA sequence investigation of Pgm found extensive cryptic amino acid polymorphism segregating with the allozyme alleles. In this study, we characterize the geographic variation of Pgm amino acid polymorphisms at the nucleotide level along a latitudinal cline in the eastern United States. A survey of 15 SNPs across the Pgm gene finds significant clinal differentiation for the allozyme polymorphisms as well as for many of the cryptic amino acid polymorphisms. A test of independence shows that pervasive linkage disequilibrium across this gene region can explain many of the amino acid clines. A single Pgm haplotype defined by two amino acid polymorphisms shows the strongest correlation with latitude and the steepest change in allele frequency across the cline. We propose that clinal selection at Pgm may in part explain the extensive amino acid polymorphism at this locus and is consistent with a multilocus response to selection in the glycolytic pathway.     

The impact of Pleistocene glaciation across the range of a widespread European coastal species

There is a growing consensus that much of the contemporary phylogeography of northern hemisphere coastal taxa reflects the impact of Pleistocene glaciation, when glaciers covered much of the coastline at higher latitudes and sea levels dropped by as much as 150 m. The genetic signature of postglacial recolonization has been detected in many marine species, but the effects of coastal glaciation are not ubiquitous, leading to suggestions that species may intrinsically differ in their ability to respond to the environmental change associated with glacial cycles. Such variation may indeed have a biological basis, but apparent differences in population structure among taxa may also stem from our heavy reliance on individual mitochondrial loci, which are strongly influenced by stochasticity during coalescence. We investigated the contemporary population genetics of Syngnathus typhle, one of the most widespread European coastal fish species, using a multilocus data set to investigate the influence of Pleistocene glaciation and reduced sea levels on its phylogeography. A strong signal of postglacial recolonization was detected at both the northern and eastern ends of the species’ distribution, while southern populations appear to have been relatively unaffected by the last glacial cycle. Patterns of population variation and differentiation at nuclear and mitochondrial loci differ significantly, but simulations indicate that these differences can be explained by the stochastic nature of the coalescent process. These results demonstrate the strength of a multilocus approach to phylogeography and suggest that an overdependence on mitochondrial loci may provide a misleading picture of population‐level processes.     

Flavonoid variation in some North Americna Saxifraga species

Eight species of Saxifraga representing sections Micranthes, Hirculus, Dactyloides and Xanthizoon were studied for their flavonoids (S. california, S. integrifolia, S. michauxii, S. ferruginea, S. eschscholtzii, S. hirculus, S. caespitosa and S. aizoides). The major compounds present in most species were kaempferol and quercetin monogluocosides and galactosides. Glucosides of kaempferol and quercetin predominate in the first four species listed, while galactosides of quercetin and myricetin are dominant in the lastthree. 3-O-Methyl- and 3,3′-di-O-methylquercetin were identified from S. californica and S. integrifolia. Saxifraga caespitosa synthesizes a complex mixture of O-methylated flavonols as well as a novel O-methylated dihydrokaempferol.Species pairs S. michauxii/S. ferruginea and S. californica/S.integrifolia exhibit close flavonoid similarity which may reflect their morphological similarities.     

Phenotypic plasticity and variation in morphological and life-history traits of antlion adults across a climatic gradient

We report here on two complementary experiments examining the effect of climate on morphological and life-history traits of antlion adults. We first examined whether body size and wing loading of emerging adults are plastic by raising larvae, collected from five antlion populations along Israel's sharp climatic gradient, in two environmental chambers simulating temperature and humidity of desert and Mediterranean climates. The variance in adult morphology was mostly related to body size, with adults of Mediterranean populations being larger than those of desert populations. Wing-to-thorax ratio was negatively correlated with temperature, compensating for the decrease in wing-beat frequency in colder environments. Differences between climatic treatments were significant for body size but not for the wing-to-thorax ratio, suggesting that body size is more plastic than the ratio between different body components. We next investigated how the exposure of antlion pupae to different climatic conditions influences the emerging adults. Adult body mass increased with final larval body mass at a faster rate when exposed to Mediterranean rather than desert conditions. Duration of the pupa stage was positively correlated with final larval mass, but only under Mediterranean conditions. Adult survival increased with initial mass (after eclosion), but was lower under desert conditions. Similarly, adults lost mass at a faster rate when exposed to desert conditions. Notably, the exposure of the pupae to varying climatic conditions had no effect on adult morphology. Climate is a major factor affecting insect life span and body size. Since body size is strongly linked to fecundity and survival, climate thus has a twofold effect on fitness: directly, and indirectly through body size.     

Spatial-temporal population dynamics across species range: from centre to margin

Understanding the boundaries of species' ranges and the variations in population dynamics from the centre to margin of a species' range is critical. This study simulated spatial-temporal patterns of birth and death rates and migration across a species' range in different seasons. Our results demonstrated the importance of dispersal and migration in altering birth and death rates, balancing source and sink habitats, and governing expansion or contraction of species' ranges in changing environments. We also showed that the multiple equilibria of metapopulations across a species' range could be easily broken following climatic changes or physical disturbances either local or regional. Although we refer to our models as describing the population dynamics across whole species' range, they should also apply to small-scale habitats (metapopulations) in which species abundance follows a humped pattern or to any ecosystem or landscape where strong central-marginal (C-M) environmental gradients exist. Conservation of both central and marginal populations would therefore be equally important considerations in making management decisions.     

Clinal variation in phyB2, a candidate gene for day-length-induced growth cessation and bud set, across a latitudinal gradient in European aspen (Populus tremula)

The initiation of growth cessation and dormancy represents a critical ecological and evolutionary trade-off between survival and growth in most forest trees. The most important environmental cue regulating the initiation of dormancy is a shortening of the photoperiod and phytochrome genes have been implicated in short-day-induced bud set and growth cessation in Populus. We characterized patterns of DNA sequence variation at the putative candidate gene phyB2 in 4 populations of European aspen (Populus tremula) and scored single-nucleotide polymorphisms in an additional 12 populations collected along a latitudinal gradient in Sweden. We also measured bud set from a subset of these trees in a growth chamber experiment. Buds set showed significant clinal variation with latitude, explaining approximately 90% of the population variation in bud set. A sliding-window scan of phyB2 identified six putative regions with enhanced population differentiation and four SNPs showed significant clinal variation. The clinal variation at individual SNPs is suggestive of an adaptive response in phyB2 to local photoperiodic conditions. Three of four SNPs showing clinal variation were located in regions with excessive genetic differentiation, demonstrating that searching for regions of high genetic differentiation can be useful for identifying sites putatively involved in local adaptation.