Genetically based population variation in aphid association with ants and predators

A species’ genotype can have extended consequences for the structure of the surrounding community, but few studies have investigated the extended consequences of genetic variation in animals. Accordingly, I examined the importance of genetically based variation among five populations of the ant-tended aphid Aphis asclepiadis for its interactions with both ants and predators. In a common environment, aphid source population accounted for 23 and 17% of the variation in the occurrence of ants and predators, respectively. Ant exclusion increased predator abundance, accounting for 25% of variation, but there was no detectable influence of ants on aphid abundance. There was an indication that aphid source populations varied in honeydew quality, but this was uncorrelated with rates of ant attendance. This study provides the first evidence for genetic variation in aphids for attractiveness to ants, and underscores the important link between intra-specific genetic variation in aphids and the processes governing arthropod community structure.     

Climatic zonation drives latitudinal variation in speciation mechanisms

Many groups of organisms show greater species richness in the tropics than in the temperate zone, particularly in tropical montane regions. Forty years ago, Janzen suggested that more limited temperature seasonality in the tropics leads to greater climatic zonation and more climatic barriers to organismal dispersal along elevational gradients in the tropics relative to temperate regions. These factors could lead to differences in how species arise in tropical versus temperate regions and possibly contribute to greater tropical diversity. However, no studies have compared the relationships among climate, elevational distribution and speciation in a group inhabiting both tropical and temperate regions. Here, we compare elevational and climatic divergence among 30 sister-species pairs (14 tropical, 16 temperate) within a single family of salamanders (Plethodontidae) that reaches its greatest species richness in montane Mesoamerica. In support of Janzen's hypothesis, we find that sister species are more elevationally and climatically divergent in the tropics than in the temperate zone. This pattern seemingly reflects regional variation in the role of climate in speciation, with niche conservatism predominating in the temperate zone and niche divergence in the tropics. Our study demonstrates how latitudinal differences in elevational climatic zonation may increase opportunities for geographical isolation, speciation and the associated build-up of species diversity in the tropics relative to the temperate zone.     

Seasonal variation in the secondary chemistry of foliar and reproductive tissues of Delphinium nuttallianum

Plant secondary compounds are critical in affecting interactions between plants and their herbivores. The norditerpene alkaloids are secondary compounds in Delphinium (larkspur) species which are divided into two classes: the N-(methylsuccinimido) anthranoyllycoctonine (MSAL-type) and non MSAL-type, and are known to be toxic to herbivorous insects and livestock. Alkaloid concentrations were measured in a whole plant context in vegetative and reproductive tissues in Delphinium nuttallianum at different stages of plant maturity at two locations to explore how plant maturity affected alkaloid concentrations within a growing season. Alkaloid concentrations differed between vegetative and reproductive tissues, with vegetative tissues having significantly lower alkaloid concentrations than reproductive tissues. However, no systematic differences in alkaloid concentrations were observed at different plant maturity stages across the growing season. Based on the data we suggest that alkaloid allocation in different plant parts of D. nuttallianum is influenced by life history of the plant, consistent with plant defense theory. At one location, as pods mature the qualitative alkaloid composition changed through structural diversification of the alkaloids present. The ecological significance of this structural diversification awaits further exploration.     

Leaf traits and performance vary with plant age and water availability in Artemisia californica

Background and AimsLeaf functional traits are strongly tied to growth strategies and ecological processes across species, but few efforts have linked intraspecific trait variation to performance across ontogenetic and environmental gradients. Plants are believed to shift towards more resource-conservative traits in stressful environments and as they age. However, uncertainty as to how intraspecific trait variation aligns with plant age and performance in the context of environmental variation may limit our ability to use traits to infer ecological processes at larger scales.MethodsWe measured leaf physiological and morphological traits, canopy volume and flowering effort for Artemisia californica (California sagebrush), a dominant shrub species in the coastal sage scrub community, under conditions of 50, 100 and 150 % ambient precipitation for 3 years.Key ResultsPlant age was a stronger driver of variation in traits and performance than water availability. Older plants demonstrated trait values consistent with a more conservative resource-use strategy, and trait values were less sensitive to drought. Several trait correlations were consistent across years and treatments; for example, plants with high photosynthetic rates tended to have high stomatal conductance, leaf nitrogen concentration and light-use efficiency. However, the trade-off between leaf construction and leaf nitrogen evident in older plants was absent for first-year plants. While few traits correlated with plant growth and flowering effort, we observed a positive correlation between leaf mass per area and performance in some groups of older plants.ConclusionsOverall, our results suggest that trait sensitivity to the environment is most visible during earlier stages of development, after which intraspecific trait variation and relationships may stabilize. While plant age plays a major role in intraspecific trait variation and sensitivity (and thus trait-based inferences), the direct influence of environment on growth and fecundity is just as critical to predicting plant performance in a changing environment.     

Climatic drivers of latitudinal variation in Late Triassic tetrapod diversity

The latitudinal biodiversity gradient (LBG), the increase in biodiversity from the poles to the equator, is one of the most widely recognized global macroecological patterns, yet its deep time evolution and drivers remain uncertain. The Late Triassic (237–201 Ma), a critical interval for the early evolution and radiation of modern tetrapod groups (e.g. crocodylomorphs, dinosaurs, mammaliamorphs), offers a unique opportunity to explore the palaeolatitudinal patterns of tetrapod diversity since it is extensively sampled spatially when compared with other pre‐Cenozoic intervals, particularly at lower palaeolatitudes. Here, we explore palaeolatitudinal patterns of Late Triassic tetrapod diversity by applying sampling standardization to comprehensive occurrence data from the Paleobiology Database (PBDB). We then use palaeoclimatic model simulations to explore the palaeoclimatic ranges occupied by major tetrapod groups, allowing insight into the influence of palaeoclimate on the palaeolatitudinal distribution of these groups. Our results show that Late Triassic tetrapods generally do not conform to a modern‐type LBG; instead, sampling‐standardized species richness is highest at mid‐palaeolatitudes. In contrast, the richness of pseudosuchians (crocodylians and their relatives) is highest at the palaeoequator, a pattern that is retained throughout their subsequent evolutionary history. Pseudosuchians generally occupied a more restricted range of palaeoclimatic conditions than other tetrapod groups, a condition analogous to modern day reptilian ectotherms, while avemetatarsalians (the archosaur group containing dinosaurs and pterosaurs) exhibit comparatively wider ranges, which is more similar to modern endotherms, such as birds and mammals, suggesting important implications for the evolution of thermal physiology in dinosaurs.     

Adaptive latitudinal variation in Common Blackbird Turdus merula nest characteristics

Nest construction is taxonomically widespread, yet our understanding of adaptive intraspecific variation in nest design remains poor. Nest characteristics are expected to vary adaptively in response to predictable variation in spring temperatures over large spatial scales, yet such variation in nest design remains largely overlooked, particularly amongst open‐cup‐nesting birds. Here, we systematically examined the effects of latitudinal variation in spring temperatures and precipitation on the morphology, volume, composition, and insulatory properties of open‐cup‐nesting Common Blackbirds’ Turdus merula nests to test the hypothesis that birds living in cooler environments at more northerly latitudes would build better insulated nests than conspecifics living in warmer environments at more southerly latitudes. As spring temperatures increased with decreasing latitude, the external diameter of nests decreased. However, as nest wall thickness also decreased, there was no variation in the diameter of the internal nest cups. Only the mass of dry grasses within nests decreased with warmer temperatures at lower latitudes. The insulatory properties of nests declined with warmer temperatures at lower latitudes and nests containing greater amounts of dry grasses had higher insulatory properties. The insulatory properties of nests decreased with warmer temperatures at lower latitudes, via changes in morphology (wall thickness) and composition (dry grasses). Meanwhile, spring precipitation did not vary with latitude, and none of the nest characteristics varied with spring precipitation. This suggests that Common Blackbirds nesting at higher latitudes were building nests with thicker walls in order to counteract the cooler temperatures. We have provided evidence that the nest construction behavior of open‐cup‐nesting birds systematically varies in response to large‐scale spatial variation in spring temperatures.     

Seasonal variation in leaf chemistry of the coast live oak Quercus agrifolia and implications for the California oak moth Phryganidia californica

Summary The perennial foliage of the California coast live oak (Quercus agrifolia Nee) permits herbivores to feed on this oak species throughout the year. Patterns of herbivory for a two-year period on Q. agrifolia were observed in relation to seasonal and age-related changes in the nutritional and defensive characteristics of leaves. Nitrogen and phosphorus contents were higher in new leaves compared to mature foliage. Structural compounds (e.g., cellulose) in leaves rapidly increased with age. Concentrations of tatal phenolics (Folin-Denis) and astringency were higher in new foliage, and concentrations of condensed tannins gradually increased as the leaves matured. Peaks of herbivore damage were observed in June and in September–October, and were caused by outbreaks of the California oak moth (Phryganidia californica). P. californica, a bivoltine oak specialist, exhibited feeding preferences in June for old leaves over emerging leaves, and showed no preferences for leaf classes in September. These results suggest that P. californica is adapted to survive on nutritionally poor foliage and to circumvent quantitative defenses such as condensed tannins.     

Thermoregulatory variation among populations of bats along a latitudinal gradient

Most studies of hibernation physiology sample individuals from populations within a single geographic area, yet some species have large ranges meaning populations likely experience area-specific levels of energetic challenges. As well, few studies have assessed within-season variation. Since physiological adjustments often are influenced by environmental factors, and the types of environments vary with geography, we expected variance in hibernation patterns among geographically separated populations. Our specific goal was to measure intraspecific variation in torpid metabolic rate (TMR) and body temperature (T _b) as a function of ambient temperature (T _a) for a non-migratory and migratory species to determine whether there is a continuum in physiological responses based on latitude. We chose big brown (Eptesicus fuscus) and eastern red bats (Lasiurus borealis) as model species and sampled individuals from populations throughout each species’ winter range. In both species, individuals from southern populations maintained higher TMR at cooler T _as and lower TMR at warmer T _as than those from northern populations. Big brown bats from southern populations regulated T _b during torpor at higher levels and there was no significant difference in T _b between populations of eastern red bats. Although metabolic responses were similar across the gradient between species, the effect was more dramatic in big brown bats. Our data demonstrate a continuum in thermoregulatory response, ranging from classic hibernation in northern populations to a pattern more akin to daily torpor in southern populations. Our research highlights the potential usefulness of bats as model organisms to address questions about within-species physiological variation in wild populations.     

Phenotypic variation and spatial structure of secondary chemistry in a natural population of a tropical tree species

To understand herbivore selection in natural plant populations, it is important to understand the landscape of plant chemical phenotypes that herbivores face and the sources of variation that will define this landscape. We studied the spatial patterns of variation in leaf secondary chemistry of the tropical tree Quararibea asterolepis , Pitt. (Bombacaceae) in a natural population on Barro Colorado Island, Panama, and used this background to discuss hypotheses of natural selection by herbivores. Quararibea plants collected from different sites had consistent differences in their chemical phenotypes. Some of these differences were explained by developmental and environmental sources of variation. Canopy trees had 13% lower yield of leaf extracts than gap seedlings, explained by 41% lower concentrations of the more abundant metabolites in the secondary compound profile. Also, plants growing in gaps had 25% higher yield than those in the understory, explained by two-fold increases in the concentration of some of the less polar secondary compounds in the profile. Differences in soil type did not affect the secondary chemistry of leaves, but sites with different topography had differences in the secondary compound profile that were not explained by any of the measured environmental sources of variation. Neighboring parent-offspring pairs and sibling/half sibling clusters displayed equal or higher variance among themselves than unrelated individuals at farther distances. Assuming that related plants should be more similar in their phenotypes, this pattern is consistent with local selection by herbivores overriding the similarity of related plants in a frequency- or distance-dependent manner.     

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.     

Comparative population genomics of latitudinal variation in Drosophila simulans and Drosophila melanogaster

Examples of clinal variation in phenotypes and genotypes across latitudinal transects have served as important models for understanding how spatially varying selection and demographic forces shape variation within species. Here, we examine the selective and demographic contributions to latitudinal variation through the largest comparative genomic study to date of Drosophila simulans and Drosophila melanogaster, with genomic sequence data from 382 individual fruit flies, collected across a spatial transect of 19 degrees latitude and at multiple time points over 2 years. Consistent with phenotypic studies, we find less clinal variation in D. simulans than D. melanogaster, particularly for the autosomes. Moreover, we find that clinally varying loci in D. simulans are less stable over multiple years than comparable clines in D. melanogaster. D. simulans shows a significantly weaker pattern of isolation by distance than D. melanogaster and we find evidence for a stronger contribution of migration to D. simulans population genetic structure. While population bottlenecks and migration can plausibly explain the differences in stability of clinal variation between the two species, we also observe a significant enrichment of shared clinal genes, suggesting that the selective forces associated with climate are acting on the same genes and phenotypes in D. simulans and D. melanogaster.     

Ineffectiveness of Two Annual Legumes as Nurse Plants for Establishment of Artemisia californica in Coastal Sage Scrub

Annual legumes are often used as nurse plants for restoration projects, but two commonly used legume species were competitors at all densities with Artemisia californica (California sagebrush), a dominant shrub of southern California coastal sage scrub. Survival of Artemisia was not reduced by the lowest densities of the native Lupinus succulentus (arroyo lupine) at ratios of Artemisia to Lupinus of 1:1 or 1:3 or by the exotic Trifolium hirtum (rose clover) at the 1:1 density, but its survival was as low as 4% at the highest densities of Trifolium (1:16) and 1:32). Overall, Trifolium was more detrimental to survival of Artemisia, but the biomass of Artemisia was reduced by 90% or more in mixtures with both legumes even at the lowest densities of 1:1. The total soil nitrogen either did not change or decreased in two of the mixtures between planting and harvest dates, indicating that the legumes not only did not add nitrogen to the soil within one growing season but even depleted it in these two cases. Whereas Lupinus had greater aboveground bio-mass than Trifolium, it had a lower root density than Trifolium. The Artemisia root system was more shallow than either Trifolium or Lupinus, possibly explaining the poor growth of Artemisia in mixtures, The legumes were one to two orders of magnitude greater in aboveground biomass than Artemisia at the 1:1 ratio and therefore may be inappropriate choices as nurse plants. There is no evidence from this study that either of these legumes can act as nurse plants, even at the lowest ratio of one nurse plant to on shrub. Nurse plants are probably more important in harsher environments than in coastal sage scrub.     

Polymorphism and geographic variation of flower color in chilean populations ofEschscholzia californica

In twelve populations ofEschscholzia californica from central Chile, where it was introduced in the latter part of the 19th century, there is a regular gradient in frequency of the allelesJ andj, which control orange vs. yellow flowers, ranging from a maximum value of 0.531 to a minimum of 0.160 forJ. Populations having higher values ofJ occupy more mesic sites. In addition, the great majority of the populations have significant excess of heterozygotes over those expected on the basis of theHardy-Weinberg equilibrium, and so exhibit balanced polymorphism for theJ-j pair of alleles, probably associated with heterosis ofJj genotypes. Evidence is presented which suggests that this locus serves as a marker for adaptative gene complexes rather than being itself associated with adaptation. In Chile, the association ofJJ genotypes with cooler, moister habitats is the opposite of the condition existing in California, and suggests that extensive reorganization of the gene pool, through changed linkage relationships, took place asE. californica populations were being established in Chile.     

Chemotypic variation of essential oils in the medicinal plant, Anemopsis californica

Anemopsis californica (Saururaceae) commonly called yerba mansa, is an important medicinal plant in many deserts in the southwestern region of North America. Populations of A. californica, collected throughout New Mexico, were examined for chemical variability in roots and rhizomes for select monocyclic (cymene, limonene, piperitone and thymol) and bicyclic (alpha-pinene, 1,8-cineole and myrtenol) monoterpenoid and phenylpropanoid (methyleugenol, isoeugenol and elemicin) derived essential oil components. Three distinct chemotypes were detected using a hierarchical clustering analysis on the concentration of 10 different analytes in three individuals from each of 17 populations. One chemotype was characterized by high elemicin concentrations, a second chemotype by high methyleugenol concentrations and the third by high piperitone and thymol concentrations. Steam distilled oil was used to screen for anticancer bioactivity. A. californica root oils demonstrated anti-proliferative activity against AN3CA and HeLa cells in vitro but no activity against lung, breast, prostate or colon cancer cells. The IC(50) values for the root oil were 0.056% and 0.052% (v/v) for the AN3CA and HeLa cells, respectively.     

Elevational parallels of latitudinal variation in the proportion of lianas in woody floras

Aim  To determine if elevational variation in the proportion of lianas in woody floras parallels the variation observed on latitudinal gradients. This is to be expected if the poleward decrease in the importance of lianas is related to the vulnerability of their wide vessels to freeze embolism.
Location  Coastal ranges of south-central Chile (latitude 37°–40° S) and western South Island of New Zealand (41°–43° S).
Methods  The presence of all woody species was recorded in plots of 2500 m² (Chile) or 100–400 m² (New Zealand) on four elevational gradients in temperate rain forest. Each species was classified as a tree, shrub or liana. Original data were obtained from 22 plots at two sites in Chile. In New Zealand, two surveys comprising a total of 296 plots were extracted from the National Vegetation Survey data base.
Results  Liana species richness declined more or less monotonically on all four gradients, whereas richness of trees and shrubs showed more varied elevational patterns. The proportion of woody species contributed by the liana life-form was negatively correlated with elevation on all four gradients, falling from 15 to 35% of the woody flora at c . 200 m a.s.l. to nil well below the tree line. The elevational and latitudinal limits of liana species were marginally significantly correlated in Chile, but not in New Zealand.
Main conclusions  The elevational parallel of the well-documented decline in liana representation with increasing latitude is consistent with the hypothesis that cold intolerance is a strong control on the global distribution of the liana life-form.     

Genetically determined variation in the azygos vein in the mouse.

The normal mouse is expected to have a single and left-sided azygos vein that develops from the paired embryonic cardinal venous system and drains most of the right and left thoracic walls into the left anterior vena cava. During routine autopsies of adult mice, most individuals of the C57BL/6J strain were found to have this pattern but a distribution of different azygos venous patterns was found in the WB/ReJ strain. In WB/ReJ the patterns varied from a single unpaired vein on the right side that connected to the right anterior vena cava through bilaterally symmetrical and paired veins to the expected unpaired vein on the left side. A classification scheme for the observed patterns of azygos veins was developed and the frequency distributions of C57BL/6J and WB/ReJ mice in these classes were compared. The strain difference in the azygos venous system between C57BL/6J and WB/ReJ can be interpreted as a genetically determined threshold trait of development. Beginning with a paired and symmetrical cardinal venous system, the C57BL/6J genotype shifts to a left-sided azygos pattern but the WB/ReJ genotype remains with a more bilateral azygos pattern. Genetic study of this azygos trait will be useful for the study of lateral asymmetries in mammalian development and for the interpretation of venous heterotaxies (anomalous placement of veins) in the mouse that are found in association with mutations such as situs inversus viscerum (iv) and dominant hemimelia (Dh).     

Integrating life history and physiology to understand latitudinal size variation in a damselfly

Our understanding of latitudinal life history patterns may benefit by jointly considering age and mass at maturity and growth rate. Additional insight may be gained by exploring potential constraints through pushing growth rates to their maximum and scoring physiological cost‐related variables. Therefore, we reared animals of a univoltine Spanish and Belgian population and of a semivoltine Swedish population of the damselfly Enallagma cyathigerum (spanning a latitude gradient of ca 2350 km) in a common environment from the eggs until adult emergence and exposed them to a transient starvation period to induce compensatory growth. Besides age and mass at maturity and growth rate we also scored investment in energy storage (i.e. triglycerides) and immune function (i.e. total activity of phenoloxidase). At emergence, body mass was greater in Spain and Sweden and lower in Belgium, suggesting a genetic component for the U‐shaped latitudinal pattern that was found also in a previous study based on field‐collected adults. The mass difference between univoltine populations can be explained by the shorter development time in the Belgian population, and this despite a higher growth rate, a pattern consistent with undercompensating countergradient variation. In line with the assumed shorter growth seasons, Belgian and Swedish animals showed higher routine growth rates and compensatory growth after transient starvation. Despite a strong link with metabolic rates (as measured by oxygen consumption) populations with higher routine growth rates had no lower fat content and had higher immune function (i.e. immune function decreased from Sweden to Spain), which was unexpected. Rapid compensatory growth did, however, result in a lowered immune function. This may contribute to the absence of perfect compensating countergradient variation in the Belgian population and the lowest routine growth rates in the Spanish population. Our results underscore the importance of integrating key life historical with physiological traits for understanding latitudinal population differentiation.     

Effects of body-size variation on flight-related traits in latitudinal populations of Drosophila melanogaster

In the present study, we tested the hypothesis whether flight-related traits such as wing area, flight-muscle ratio, wing loading and dispersal yield evidence of geographical variation in nine wild-collected as well as laboratory-reared (at 21°C) latitudinal populations of Drosophila melanogaster from the Indian subcontinent. We observed positive clinal variation in the wing–thorax ratio, wing aspect ratio and wing area, along a latitudinal gradient for both the sexes. In contrast, geographical changes in three parameters of flight ability, i.e. flight-muscle ratio, wing loading and dispersal, showed negative correlation withlatitude. On the basis of isofemale line variability, we observed positive correlation of wing loading with flight-muscle ratio as well as dispersal behaviour in both the sexes. We also found positive correlation between duration of development and wing area. Interestingly, southern populations of D. melanogaster from warm and humid habitats exhibited higher flight-muscle ratio as well as the higher wing loading than northern populations which occur in cooler and drier climatic conditions. Laboratory tests for dispersal-related walking behaviour showed significantly higher values for southern populations compared with northern populations of D. melanogaster. Multiple regression analysis of geographical changes in flight-muscle ratio, wing loading as well as walking behaviour as a function of average temperature and relative humidity of the origin of populations in wild-collected flies have suggested adaptive changes in flight-related traits in response to steeper gradients of climatic factors in the Indian subcontinent. Finally, adaptive latitudinal variations in flight-related traits in D. melanogaster are consistent with results of other studies from different continents despite differences due to specific climatic conditions in the Indian subontinent.