Patterns Of Variation In Traits Research Articles

Genetics of body shape and armour variation in threespine sticklebacks

Patterns of genetic variation and covariation can influence the rate and direction of phenotypic evolution. We explored the possibility that the parallel morphological evolution seen in threespine stickleback (Gasterosteus aculeatus) populations colonizing freshwater environments is facilitated by patterns of genetic variation and covariation in the ancestral (marine) population. We estimated the genetic (G) and phenotypic (P) covariance matrices and directions of maximum additive genetic (g(max) ) and phenotypic (p(max) ) covariances of body shape and armour traits. Our results suggest a role for the ancestral G in explaining parallel morphological evolution in freshwater populations. We also found evidence of genetic constraints owing to the lack of variance in the ancestral G. Furthermore, strong genetic covariances and correlations among traits revealed that selective factors responsible for threespine stickleback body shape and armour divergence may be difficult to disentangle. The directions of g(max) and p(max) were correlated, but the correlations were not high enough to imply that phenotypic patterns of trait variation and covariation within populations are very informative of underlying genetic patterns.

Differentiation in a tropical deceptive orchid: colour polymorphism and beyond

Colour polymorphism has often been described among individuals of deceptive orchid species, and several studies have investigated reproductive success variations among colour morphs. However, whether colour morphs differed in other traits has received little attention in previous studies. Here, we report the case of a tropical deceptive orchid in Reunion Island that exhibits three different colour varieties. We investigated patterns of trait variation among colour varieties and found significant differences in floral and inflorescence morphology. Interestingly, we found that most populations included only individuals of a single variety, and that strong differences exist among varieties in spatial distribution and flowering phenology. This situation differs from previously reported cases of floral polymorphism in deceptive orchids where colour morphs co-occur and flower in the same populations. The spatio-temporal variation in flowering suggests that colour varieties have independent reproductive ecologies and are adapted to local conditions. We propose that potential variations in pollinator species abundance or diversity and the co-occurrence of nectar-producing species in the community may have driven the adaptation of each variety to its current pollination niche.

Concordance between locomotor morphology and foraging mode in lacertid lizards

Foraging behaviors exist along a continuum from highly sedentary, ambush foraging, to more continuous searching, or active foraging. Foraging strategies, or modes, are defined based upon locomotor behaviors (e.g. percent time moving, moves per minute). In lizards, traits correlated with ambush and active foraging have been of interest for some time; however, general patterns of correlated evolution between locomotor morphology and locomotor behavior have only recently begun to be quantified. In this study, variation in hindlimb morphology is investigated in a model group of lizard species that vary between active foraging and more sedentary (or mixed) foraging mode. Canonical variates analysis reveals that the two active foraging species occupy similar regions of the morphospace, while the two more sedentary species occupy different regions. The active foraging species have a narrow pelvis with shorter tibia and femora. The more sedentary species have a wide pelvis, long tibia and femora, and slightly longer metatarsals. Phylogenetic patterns of trait variation were examined through ancestral character state reconstruction and show morphological shifts in concert with foraging mode in these species. The observed shifts in locomotor morphology are discussed in light of published data on sprint speed and endurance in these species. Together, the data show that linking morphological variation to variation in stride length and stride frequency is critical to understanding the evolution of locomotor performance. Much more stride length and frequency data are needed among ambush, mixed, and active foraging species because these parameters, and their morphological components, are likely correlated with variation in food acquisition mode.

Establishment of parallel altitudinal clines in traits of native and introduced forbs

Due to altered ecological and evolutionary contexts, we might expect the responses of alien plants to environmental gradients, as revealed through patterns of trait variation, to differ from those of the same species in their native range. In particular, the spread of alien plant species along such gradients might be limited by their ability to establish clinal patterns of trait variation. We investigated trends in growth and reproductive traits in natural populations of eight invasive Asteraceae forbs along altitudinal gradients in their native and introduced ranges (Valais, Switzerland, and Wallowa Mountains, Oregon, USA). Plants showed similar responses to altitude in both ranges, being generally smaller and having fewer inflorescences but larger seeds at higher altitudes. However, these trends were modified by region-specific effects that were independent of species status (native or introduced), suggesting that any differential performance of alien species in the introduced range cannot be interpreted without a fully reciprocal approach to test the basis of these differences. Furthermore, we found differences in patterns of resource allocation to capitula among species in the native and the introduced areas. These suggest that the mechanisms underlying trait variation, for example, increasing seed size with altitude, might differ between ranges. The rapid establishment of clinal patterns of trait variation in the new range indicates that the need to respond to altitudinal gradients, possibly by local adaptation, has not limited the ability of these species to invade mountain regions. Studies are now needed to test the underlying mechanisms of altitudinal clines in traits of alien species.

Disentangling composite colour patterns in a poison frog species

A phylogenetic approach was performed to infer whether variation in conspicuous colour-patterns of a poison frog (Dendrobatidae: Dendrobates tinctorius) has evolved neutrally or under selection. Colour and pattern were split into components that were separately analysed and subsequently re-grouped via principal component analysis. This revealed four different ‘displayed’ factors on the dorsal and lateral views versus one ‘concealed’ factor on the ventral view. Based on the assumption that current patterns of trait variation contain information about the evolutionary history of the phenotype, we correlated these trait components to a neutrally evolving gene fragment (cytochrome b). The concealed factor was significantly correlated with the marker fragment, which identified it as having evolved under genetic drift. Noncorrelation of all displayed factors with the marker may indicate the evolution of colour patterns on dorsum and flanks under selection. In our example, colour pattern should therefore be regarded as a multicomponent signal system.

Evolutionary responses of discontinuous gas exchange in insects.

The discontinuous gas-exchange cycles (DGCs) observed in many quiescent insects have been a cause of debate for decades, but no consensus on their evolutionary origin or adaptive significance has been achieved. Nevertheless, three main adaptive hypotheses have emerged: (i) the hygric hypothesis suggests that DGCs reduce respiratory water loss; (ii) the chthonic hypothesis suggests that DGCs facilitate gas exchange during environmental hypoxia, hypercapnia, or both; and (iii) the oxidative-damage hypothesis suggests that DGCs minimize oxidative tissue damage. However, most work conducted to date has been based on single-species investigations or nonphylogenetic comparative analyses of few species, despite calls for a strong-inference, phylogenetic approach. Here, we adopt such an approach by using 76 measurements of 40 wild-caught species to examine macrophysiological variation in DGC duration in insects. Potential patterns of trait variation are first identified on the basis of the explicit a priori predictions of each hypothesis, and the best phylogenetic generalized least-squares fit of the candidate models to the data is selected on the basis of Akaike's information criterion. We find a significant positive relationship between DGC duration and habitat temperature and an important interaction between habitat temperature and precipitation. This result supports the hygric hypothesis. We conclude that the DGCs of insects reduce respiratory water loss while ensuring adequate gas exchange.

Biological traits of benthic macroinvertebrates in California mediterranean-climate streams: long-term annual variability and trait diversity patterns

We examined long-term variability of the biological traits of benthic macroinvertebrate communities using data collected annually (20-, 7-, and 8-years in duration, respectively) from three study regions in northern California's mediterranean-climate region. We related temporal variability to several climatic and physical variables representing precipitation and hydrologic conditions, and assessed the stability of biological-trait composition using time-lag regression analysis. We also examined the temporal variability of trait diversity metrics and their relationship to taxa richness. Trait modalities exhibited high temporal variability (∼ 20 % of all trait modalities had a CV > 50 %) and biological-trait composition varied along a gradient from dry (extreme drought) to wet (above average) years. Traits conferring resistance or resilience to drying (e.g. desiccation resistance, aerial respiration) were more common in drought years, whereas traits conferring resistance or resilience to floods (e.g. flattened body shape, drift dispersal) were more common in wet years. Furthermore, prolonged drought and the patterns of increasing or decreasing precipitation over time in the study regions resulted in patterns of directional change in biological-trait composition. Although biological traits had high temporal variation within each region, between-region variation was higher than temporal variation in a single region. As found in other studies, trait diversity and richness metrics were relatively invariable over time despite high taxonomic turnover. However, there was considerable regional variation in trait diversity metrics along a gradient of increasing taxonomic diversity. The four functional metrics studied showed high redundancy. Taxa richness was positively correlated with three functional metrics (trait richness and two trait diversity measures), and trait richness and diversity became saturated taxa richness increased (∼50 taxa). In communities largely controlled by abiotic factors, such as stream invertebrates in mediterranean-climate, high redundancy may be the result of strong filtering of traits at larger spatial scales.

How and When Selection Experiments Might Actually be Useful

Laboratory natural selection and artificial selection are vital tools for addressing specific questions about evolutionary patterns of variation. Laboratory natural selection can illuminate whether a putative selective agent is capable of generating long-term, sustained changes in individual traits and suites of traits. Artificial selection is the essential tool for understanding the general evolvability of traits and the extent to which genetic correlations constrain evolution. We review the contexts in which each type of experiment seems capable of offering key insights into important evolutionary issues. We also discuss theoretical and methodological considerations that play critical roles in designing selection experiments that are relevant to evolutionary patterns of trait variation. In particular, we focus on the critical role of selection intensity and the consequences of experiments with different intensities. While selection experiments are not practical in many cases, sophisticated selection experiments-designed with careful consideration of the theory of selection-should be taken beyond model organisms and used in well-chosen natural systems to understand natural patterns of variation.

Components of phenotypic variation in the morphometrics of shearwater (Puffinus) species

Coefficients of variation were used to examine the variability of bill, wing and foot measurements of 11 sexually size dimorphic and seven monomorphic shearwater (Puffinus) species, and to investigate the selective regimes that may be acting on these traits. The effects of phylogeny can lead to a lack of statistical independence among interspecies data, so randomization tests were performed in addition to standard statistical approaches, which assume independence. However, the results obtained from the two approaches always agreed. Among the 18 species, bill measurements were significantly more variable than those of the wing or foot; furthermore, bill depth dimensions exhibited the greatest amount of phenotypic variation. Both sexually monomorphic and sexually dimorphic species exhibited the same patterns of phenotypic variation. In the sexually dimorphic species, patterns of trait variation did not differ significantly between sexes. The findings suggest that variation in the bill of dimorphic Puffinus species is not due to sexual selection. More probably, variability among the traits is due to differences in the strength of natural selection, with those traits under strong stabilizing selection (e.g. wing, tarsus and mid‐toe) exhibiting reduced variability, as optimum physical dimensions are being selected for.

The Evolution of Plant Functional Variation: Traits, Spectra, and Strategies

Variation in plant functional traits results from evolutionary and environmental drivers that operate at a variety of different scales, which makes it a challenge to differentiate among them. In this article we describe patterns of functional trait variation and trait correlations within and among habitats in relation to several environmental and trade‐off axes. We then ask whether such patterns reflect natural selection and can be considered plant strategies. In so doing we highlight evidence that demonstrates that (1) patterns of trait variation across resource and environmental gradients (light, water, nutrients, and temperature) probably reflect adaptation, (2) plant trait variation typically involves multiple‐correlated traits that arise because of inevitable trade‐offs among traits and across levels of whole‐plant integration and that must be understood from a whole‐plant perspective, and (3) such adaptation may be globally generalizable for like conditions; i.e., the set of traits (collections of traits in syndromes) of taxa can be considered as “plant strategies.”

Variation among populations ofClarkia Unguiculata(Onagraceae) along altitudinal and latitudinal gradients

We investigated phenotypic variation in 15 traits in greenhouse-grown plants from 16 populations of Clarkia unguiculata from three elevational habitats and six latitudinal transects. Populations from the lowest and highest elevations were geographically and ecologically marginal within the species' range. We (1) describe patterns of trait variation with elevation and latitude; (2) compare latitutidinal variation between marginal and central areas of the species' range; and (3) compare patterns of variation within C. unguiculata to interspecific patterns within the genus. Although there was some evidence that traits varied clinally (i.e., increased/decreased monotonically) along environmental gradients, interaction effects between altitude and latitude dominated patterns of variation. For most traits, latitudinal trends at the low-elevation margin of the species' range differed from trends at mid- and high-elevation areas. Based on interspecific comparisons, populations at the hotter, more arid ends of both environmental gradients were expected to have rapid development, small flowers and vegetative size, low levels of herkogamy and protandry, and high rates of gas exchange. Instead, we found that while some traits were correlated with one gradient in the expected way (e.g., development time with elevation, gas-exchange physiology with latitude), all traits were not consistently associated with each other along both gradients, and intraspecific patterns of variation differed from interspecific patterns.

Seed Size and Phylogeny in Six Temperate Floras: Constraints, Niche Conservatism, and Adaptation

Seed size is correlated with a number of other plant attributes, and such correlations are commonly interpreted as adaptations to different life histories. However, seed size is also strongly related to species phylogeny. In the current comparative literature, correlations between trait variation and taxonomic membership are often interpreted as evidence for phylogenetic constraints, and such phylogenetic explanations of variation are presented as alternatives to adaptive explanations. However, phylogenetic patterns of trait variation should not be discussed exclusively in terms of constraints. Neither should phylogenetic and adaptive explanations for variation be automatically presented as alternatives. Another explanation for similarity among related taxa is phylogenetic niche conservatism (PNC). This process is well known to exist-indeed, it is an inevitable consequence of evolution by natural selection-and is explicitly adaptive. Under PNC, radiation within a lineage is shaped by phylogeny, by the env...

Genecology of Douglas‐Fir in a Watershed in the Oregon Cascades

To gain insight into genetic microstructure of subregional populations of coastal Douglas—fir (Pseudotsuga menziesii [Mirb.] Franco var. menziesii), genetic variability in a population found on a 6100—ha, central Oregon watershed is described. Genotypic values of 193 parent trees located throughout the watershed were estimated from progeny grown in a common garden. Then, genetic variation was partitioned into components attributable to parent—tree location and to differences among trees within locations. Within—location variation appeared to be homogeneous in the watershed; between—location variation was related to topography, but the patterns of trait variation differed depending on the trait measured. Growth traits exhibited strong gradients with elevation, but the relationship varied depending on position within the watershed. Based on estimates of proportions of nonadapted seedlings in hypothetical transfers of seed from one part of the watershed to another, genetic differentiation within the watershed was surprisingly large. In one transfer of 3.5 km between north— and south—facing slopes at the same elevation, °80% of seedlings were estimated to be poorly adapted. Although other possibilities exist, the topoclinal variation in traits probably results from selection as does the large within—location variation. Both kinds of variation are due to high selection intensities in the seedling stage, the former to selection by average environmental differences along gradients, the latter to microenvironmental heterogeneity. The combination of high within— and between—habitat variation is suited to a species which episodically colonizes an environment that is extremely heterogeneous in time and space.