Does intraspecific competition promote variation? A test via synthesis

Competitive diversification, that is, when increasing intraspecific competition promotes population niche expansion, is commonly invoked in evolutionary studies and currently plays a central role in how we conceptualize the process of adaptive diversification. Despite the frequency with which this idea is cited, the empirical evidence for the process is somewhat limited, and the findings of these studies have yet to be weighed objectively through synthesis. Here, we sought to fill this gap by reviewing the existing literature and collecting the data necessary to assess the evidence for competition as a diversifying force. Additionally, we sought to test a more recent hypothesis, which suggests that competition can act to both promote and inhibit dietary diversification depending on the degree to which a consumer depletes its resources. The surprising result of this synthesis was that increasing competition did not have a mean positive effect on population‐level diet breadth or the degree of individual specialization. Instead, we found that increasing intraspecific competition had a restricting effect on population‐level diet breadth in as many cases as it had a diversifying effect. This wide disparity in the effect of competition on consumer diet variation was negatively related to a metric for consumer resource depletion. Altogether, these findings call into question a long‐standing assumption of basic evolutionary models and lend some support to recent theoretical predictions. Specifically, these findings support the idea that competition is primarily diversifying for species with a small effect (per unit biomass) on their resources and that resource depletion limits the diversifying effect of competition for consumers with larger ecological effects.     

How universal are reserve design rules? A test using butterflies and their life history traits

We used butterfly species lists available for a set of 125 Czech Republic National Nature Reserves and Monuments, the highest small‐sized conservation category in the country encompassing practically all biotope types existing in central Europe, to test the validity of generally agreed “reserve design rules” using multivariate ordination analyses. Further, we used ordination analysis of butterfly life history traits to seek for biological mechanisms responsible for butterfly community responses to essentially geometric reserves characteristics. Reserve area, relative perimeter, within reserve habitat heterogeneity, and surrounding landscape compositional and configurational heterogeneity all affected the composition of butterfly assemblages after controlling for effects of geographical position and prevailing biotope type. Species inclining towards large reserves displayed low mobility and high local population density, probably because they require large habitat areas to maintain self‐sustaining populations; such species tend to have restricted distribution in the country and threatened status. Reserves with relatively long boundaries hosted species with high mobility, broad trophic range and long adult period; faunas of such reserves contain high proportions of widespread generalists. Species with narrow trophic ranges inclined towards reserves containing diverse habitats, probably due to requirements for high floristic diversity. Species with short adult flight, low generations number and overwintering in early stages inclined towards reserves situated amidst diverse landscapes, perhaps because such species require finely‐grained mosaics for metapopulation dynamics. Commonly agreed reserve design rules thus hold for Central European butterflies, but different design characteristics are important for individual species, depending on their life histories.     

The relationship between geographic range size and life history traits: is biogeographic history uncovered? A test using the Iberian butterflies

The geographic range of a species is influenced by past phylogenetic and biogeographic patterns. However, other historical interactions, including the interplay between life history and geography, are also likely involved. Therefore, the range size of a species can be explained on the basis of niche‐breadth or dispersal related hypotheses, and previous work on European butterflies suggests that both, under the respective guise of ecological specialisation and colonising ability may apply. In the present study, data from 205 species of butterflies from the Iberian peninsula were processed through multiple regression analyses to test for correlations between geographic range size, life history traits and geographic features of the species distribution types. In addition, the percentage of variance explained by the subsets of variables analyzed in the study, with and without control for phylogenetic effects was tested. Despite a complex pattern of bivariate correlations, we found that larval polyphagy was the single best correlate of range size, followed by dispersal. Models that combined both life history traits and geographic characteristics performed better than models generated independently. The combined variables explained at least 39% of the variance. Bivariate correlations between range size and body size, migratory habits or egg size primarily reflected taxonomic patterning and reciprocal correlations with larval diet breadth and adult phenology. Therefore, aspects of niche breadth i.e. potential larval diet breadth emerged as the most influential determinants of range size. However, the relationships between these types of ecological traits and biogeographic history must still be considered when associations between life history and range size are of interest.     

Do interspecific interactions between females drive shifts in habitat use? A test using the lizards Anolis carolinensis and A. sagrei

An important goal in evolutionary ecology is to understand how and why coexisting closely related species partition habitat among themselves. Although studies of interspecific interactions typically focus on males, interactions between females may also play an important role in shaping habitat use within multi‐species communities. The green anole (Anolis carolinensis) exhibits a wide range of habitat use in south‐eastern Louisiana, but its observed habitat use is restricted and altered in areas where it occurs with the introduced Anolis sagrei. We staged interactions between these two species in the laboratory to test the hypothesis that A. sagrei dominate A. carolinensis in contests over shared habitat. We examined whether species identity, bite force, dewlap size, and body size affected the outcome of interspecific interactions between both males and females, and tested the prediction that bite force and size would be the most important determinants of interaction outcomes in both sexes. In male interspecific interactions, we found that individuals with relatively larger dewlaps tended to score higher on aggressive behaviours regardless of species identity, and that interactions consisted of signalling and rarely escalated to physical combat. However, we found that A. sagrei females achieved higher aggressive scores than A. carolinensis females in almost all cases, lending support to the notion that female interspecific behaviour is probably more important than male behaviour in driving changes in habitat use. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 843–851.     

A willow drawing from 1786: the earliest depiction of intraspecific trait variation in plants?

Background and AimsThe study of intraspecific trait variation (ITV) in plants has a long history, dating back to the fourth century BC. Its existence was widely acknowledged by the end of the 18th century, although systematic and experimental studies commenced only a century later. However, the historiography of ITV has many gaps, especially with regard to early observations and visual documents. This note identifies an early depiction of plant ITV.MethodsThe botanical works of Johann Wolfgang von Goethe (1749–1832), a German writer and naturalist, were subjected to close reading. This included all publications and unpublished sources related to botany between 1785 and 1832 (e.g. notes, drafts, diaries, letters, drawings). This material is accessible in the multi-volume historical-critical edition of Goethe’s studies in natural science (Leopoldina-Ausgabe).Key ResultsA diary entry from 9 September 1786 described changes in leaf morphology along an elevation gradient in the Bavarian Alps. The leaves of an unidentified species of willow (Spix sp.) and gentian (Gentiana sp.) were said to become narrower with increasing elevation; leaves also stood further apart on twigs, and the latter became thinner. A crude drawing of two willow twigs illustrated the differences. Goethe conjectured that the differences were due to environmental conditions.ConclusionsGoethe’s notes were anecdotal, and it is unclear whether the observed plant individuals actually belonged to the same species. Nevertheless, the notes represent an early and clear articulation of the hypothesis that changes in environmental conditions can cause ITV in a natural plant population. The drawing may be the earliest visual record of environmentally caused plant ITV in the wild.     

Sequence variation patterns along a latitudinal cline in Scots pine (Pinus sylvestris): signs of clinal adaptation?

Adaptive clinal variation is abundant in nature, and its genetic basis is of great interest. The polygenic nature of this variation poses a challenge for identifying the causative loci underlying these adaptations. Here, we have examined the patterns of sequence variation in ten candidate genes for timing of bud set and cold tolerance, traits that form strong latitudinal clines in Scots pine (Pinus sylvestris). A set of reference genes was used for comparison and to infer a simplified demographic background model with approximate Bayesian computation methods. Against the resulting bottleneck model, the neutrality tests show little signs for local adaptation, but species-wide selection is suggested in some of the studied genes. In line with the theoretical expectations, we see little evidence for adaptive differentiation with F _ST methods. Instead, allele frequency clines were found in three genes (dhn1, ftl2, and prr1). Our results add to the discussion on which molecular signals best characterize a gene subject to clinally varying selection. This will be especially relevant when these kinds of observations can be examined in parallel with association study results.     

Do extreme environments provide a refuge from pathogens? A phylogenetic test using serpentine flax

Abiotically extreme environments are often associated with physiologically stressful conditions, small, low-density populations, and depauperate flora and fauna relative to more benign settings. A possible consequence of this may be that organisms that occupy these stressful habitats receive fitness benefits associated with reductions in the frequency and/or intensity of antagonistic species interactions. I investigated a particular form of this effect, formalized as the "pathogen refuge hypothesis," through a study of 13 species of wild flax that grow on stressful serpentine soils and are often infected by a pathogenic fungal rust. The host species vary in the degree of their serpentine association: some specialize on extreme serpentine soils, while others are generalists that occur on soils with a wide range of serpentine influence. Phylogenetically explicit analyses of soil chemistry and field-measured disease levels indicated that rust disease was significantly less frequent and severe in flax populations growing in more stressful, low-calcium serpentine soils. These findings may help to explain the persistence of extremophile species in habitats where stressful physical conditions often impose strong autecological fitness costs on associated organisms. Ancestral state reconstruction of serpentine soil tolerance (approximated using soil calcium concentrations) suggested that the ability to tolerate extreme serpentine soils may have evolved multiple times within the focal genus.     

Do life history traits predict responses to defoliation in co‐occurring prairie grasses?

Abstract. This study examines whether competition between the unpalatable grass Hilaria mutica and three co‐occurring, palatable grasses in a Texan mixed prairie is altered by non‐selective or selective defoliation. In this four‐year study, plants were grown in monoculture or in combination with the unpalatable Hilaria in a replacement design. Under no defoliation, the unpalatable Hilaria had a lower growth potential than Bouteloua curtipendula and Nassella leucotricha that were of equal stature, and produced only as much as the shorter grass, Buchloe dactyloides. Bouteloua had the highest growth potential under no‐defoliation and was defoliation tolerant, except when defoliated at ground level. Nassella was more productive than the unpalatable Hilaria, since the ability to grow earlier in the year enabled it to compete successfully with Hilaria. These results indicate that with adequate deferment Bouteloua and Nassella should compete successfully with Hilaria and Buchloe should be able to maintain itself in the presence of Hilaria. Under non‐selective defoliation, Hilaria was able to compete successfully only with Buchloe. Hilaria was sensitive to defoliation, despite being rhizomatous, and competed less successfully with Buchloe after non‐selective defoliation than it did when not defoliated. This indicates that the management practice of burning and stocking heavily with livestock until Hilaria is avoided, resulting in non‐selective defoliation, will not cause Hilaria to be more competitive with the more palatable Bouteloua, Buchloe or Nassella. Hilaria was able to compete most successfully under selective defoliation when it was not defoliated. Under selective defoliation, by avoiding herbivory, Hilaria is able to compete strongly with at least Buchloe and Nassella. The reaction of Nassella and Buchloe to selective defoliation indicates that they may have been displaced by Hilaria in the past. In contrast, under the short‐term and non‐limiting growth conditions of this study, Bouteloua competed successfully with Hilaria even under selective defoliation. These results do not rule out the possibility that, through selective defoliation, Hilaria may have displaced other grasses including Bouteloua in the past.     

Does environmental heterogeneity drive functional trait variation? A test in montane and alpine meadows

While community‐weighted means of plant traits have been linked to mean environmental conditions at large scales, the drivers of trait variation within communities are not well understood. Local environmental heterogeneity (such as microclimate variability), in addition to mean environmental conditions, may decrease the strength of environmental filtering and explain why communities support different amounts of trait variation. Here, we assess two hypotheses: first, that more heterogeneous local environments and second, that less extreme environments, should support a broader range of plant strategies and thus higher trait variation. We quantified drivers of trait variation across a range of environmental conditions and spatial scales ranging from sub‐meter to tens of kilometers in montane and alpine plant communities. We found that, within communities, both environmental heterogeneity and environmental means are drivers of trait variation. However, the importance of each environmental factor varied depending on the trait. Our results indicate that larger‐scale trait–climate linkages that hold across communities also apply at small spatial scales, suggesting that microclimate variation within communities is a key driver of community functional diversity. Microclimatic variation provides a potential mechanism for helping to maintain diversity in local communities and also suggests that small‐scale environmental heterogeneity should be measured as a better predictor of functional diversity.     

Do highly divergent loci reside in genomic regions affecting reproductive isolation? A test using next-generation sequence data in Timema stick insects

ABSTRACT: BACKGROUND: Genetic divergence during speciation with gene flow is heterogeneous across the genome, with some regions exhibiting stronger differentiation than others. Exceptionally differentiated regions are often assumed to experience reduced introgression, i.e., reduced flow of alleles from one population into another because such regions are affected by divergent selection or cause reproductive isolation. In contrast, the remainder of the genome can be homogenized by high introgression. Although many studies have documented variation across the genome in genetic differentiation, there are few tests of this hypothesis that explicitly quantify introgression. Here, we provide such a test using 38,304 SNPs in populations of Timema cristinae stick insects. We quantify whether loci that are highly divergent between geographically separated ('allopatric') populations exhibit unusual patterns of introgression in admixed populations. To the extent this is true, highly divergent loci between allopatric populations contribute to reproductive isolation in admixed populations. RESULTS: As predicted, we find a substantial association between locus-specific divergence between allopatric populations and locus-specific introgression in admixed populations. However, many loci depart from this relationship, sometimes strongly so. We also report evidence for selection against foreign alleles due to local adaptation. CONCLUSIONS: Loci that are strongly differentiated between allopatric populations sometimes contribute to reproductive isolation in admixed populations. However, geographic variation in selection and local adaptation, in aspects of genetic architecture (such as organization of genes, recombination rate variation, number and effect size of variants contributing to adaptation, etc.), and in stochastic evolutionary processes such as drift can cause strong differentiation of loci that do not always contribute to reproductive isolation. The results have implications for the theory of 'genomic islands of speciation'.     

What causes geographical variation in the species–area relationships? A test from forests in China

The increase of species richness with sampling area and the decrease with latitude and altitude are two of the most frequently studied patterns in biogeography. However, few studies have simultaneously examined these two patterns to investigate how species–area relationships (SAR) vary with latitude and altitude. In this study, we explore the spatial patterns of SAR in forests in China by investigating numbers of species by life form group (trees, shrubs and herbs) in 32 nested plots from 12 mountains ranging from 18.7°N to 51.9°N in latitude and from 300 to 3150 m in altitude. The slopes of the power law SAR (z‐values) decreased with increasing latitude for all life forms except herbaceous plants, and also decreased with increasing altitude for all life forms but not for shrubs. Latitude and altitude, as well as their interactions, together explained 65.4, 61.8, 48.9 and 45.3% of the variation in z‐values for overall species, trees, shrubs and herbaceous plants, respectively. In addition, actual evapotranspiration affected SAR significantly, but this effect varied significantly among life forms. We concluded that there are significant geographical patterns of SAR for China's forests, which is primarily controlled by energy availability.     

Drugs and parasites: global experiments in life history evolution?

Recent work has shown that the evolution of Drosophila melanogaster resistance to attack by the parasitoid Asobara tabida is constrained by a trade-off with larval competitive ability. However, there are two very important questions that need to be answered. First, is this a general cost, or is it parasitoid specific? Second, does a selected increase in immune response against one parasitoid species result in a correlated change in resistance to other parasitoid species? The answers to both questions will influence the coevolutionary dynamics of these species, and also may have a previously unconsidered, yet important, influence on community structure.     

Parental guidance? Trans-generational influences on offspring life history in mosquitoes

Parental effects are important factors that might influence evolutionary and ecological aspects of parasite vectors and the parasites they transmit. A recent study demonstrated the importance of parental rearing conditions on the malaria vector Anopheles stephensi. When parents are reared in a food-limited environment their offspring have increased bloodmeal sizes and larger clutches. The study highlights that ecological studies are vital for understanding vectors of disease and ultimately for developing effective control strategies.     

Do changes in DNA methylation mediate or interact with SNP variation? A pharmacoepigenetic analysis

Background

In studies with multi-omics data available, there is an opportunity to investigate interdependent mechanisms of biological causality. The GAW20 data set includes both DNA genotype and methylation measures before and after fenofibrate treatment. Using change in triglyceride (TG) levels pre- to posttreatment as outcome, we present a mediation analysis that incorporates methylation. This approach allows us to simultaneously consider a mediation hypothesis that genotype affects change in TG level by means of its effect on methylation, and an interaction hypothesis that the effect of change in methylation on change in TG levels differs by genotype. We select 322 single-nucleotide polymorphism–cytosine-phosphate-guanine (SNP-CpG) site pairs for mediation analysis on the basis of proximity and marginal genome-wide association study (GWAS) and epigenome-wide association study (EWAS) significance, and present results from the real-data sample of 407 individuals with complete genotype, methylation, TG levels, and covariate data.

Results

We identified 3 SNP-CpG site pairs with significant interaction effects at a Bonferroni-corrected significance threshold of 1.55E-4. None of the analyzed sites showed significant evidence of mediation. Power analysis by simulation showed that a sample size of at least 19,500 is needed to detect nominally significant indirect effects with true effect sizes equal to the point estimates at the locus with strongest evidence of mediation.

Conclusions

These results suggest that there is stronger evidence for interaction between genotype and methylation on change in triglycerides than for methylation mediating the effect of genotype.

     

Do aposematism and Batesian mimicry require bright colours? A test,using European viper markings.

Predator avoidance of noxious prey, aposematism and defensive mimicry are normally associated with bright, contrasting patterns and colours. However, noxious prey may be unable to evolve conspicuous coloration because of other selective constraints, such as the need to be inconspicuous to their own prey or to specialist predators. Many venomous snakes, particularly most vipers, display patterns that are apparently cryptic, but nevertheless highly characteristic, and appear to be mimicked by other, non-venomous snakes. However, predator avoidance of viper patterns has never been demonstrated experimentally. Here, the analysis of 813 avian attacks on 12,636 Plasticine snake models in the field shows that models bearing the characteristic zigzag band of the adder (Vipera berus) are attacked significantly less frequently than plain models. This suggests that predator avoidance of inconspicuously but characteristically patterned noxious prey is possible. Our findings emphasize the importance of mimicry in the ecological and morphological diversification of advanced snakes.     

Fertility and mortality patterns of captive Bornean and Sumatran orangutans: is there a species difference in life history?

Across broad taxonomic groups, life history models predict that increased ecological predictability will lead to conservative investment in reproductive effort. Within species, however, organisms are predicted to have increased reproductive rates under improved environmental conditions. It is not clear how these models apply to closely-related species. In this paper, we examine predictions from these models as applied to variability in reproductive rates between the two species of orangutans, Pongo pygmaeus (Bornean) and Pongo abelii (Sumatran). Orangutans exhibit characteristics of a "slow" life history strategy with large bodies, late age at maturity, low reproductive rates, and long lifespan. Recently, researchers proposed that Sumatran orangutans may have an even slower life history than Bornean orangutans as a result of ecological and genetic differences (Wich et al., 2004). We examined this hypothesis by studying important aspects of life history of both species under conditions of relative ecological stability, in captivity. In this large dataset, there were no significant species differences in age of first or last reproduction, completed fertility, perinatal and postnatal mortality, or female longevity. Bornean orangutans in captivity did have significantly longer interbirth intervals, and male Bornean orangutans had higher survival past maturity. Our results do not support the hypothesis that selection has led to decreased reproductive effort under conditions of increased habitat quality in Sumatra (Wich et al., 2004), and instead suggest that phenotypic flexibility may be particularly important in explaining differences between closely related species.     

Do differences in plasticity during early growth lead to differing success in competition? A test using four co‐occurring annual Papaver

Plant species differ in their ability to transform available resources to biomass and to respond in a plastic way to environmental circumstances; we hypothesized that such differences among four weed taxa of Papaver would explain differences in their competitive response. We first compared two populations each of Papaver rhoeas L., P. dubium L. ssp. dubium, P. dubium L. ssp. lecoqii (Lamotte) Syme and P. argemone L., grown in a greenhouse for 6 weeks in a nutrient gradient combined with two light treatments to elucidate possible differences in responses. As there were clear differences, a second experiment evaluated whether these differences also meant differences in competitive response, during early growth, when tested against two crops (wheat, rape). The assumption that competitive response was linked to the ability to transform nutrient and light to biomass was not supported: even though differences in extent of plasticity existed, the effect of competition was similar for the taxa. Thus, higher plasticity and ability to transform available recourses to biomass did not lead to stronger competitiveness for Papaver during early growth.