Physiological adjustments to arid and mesic environments in larks (Alaudidae)

Because deserts are characterized by low food availability, high ambient temperature extremes, and absence of drinking water, one might expect that birds that live in these conditions exhibit a lower basal metabolic rate (BMR), reduced total evaporative water loss (TEWL), and greater ability to cope with high air temperatures than their mesic counterparts. To minimize confounding effects of phylogeny, we compared the physiological performance of four species of larks at ambient temperatures (T(a)'s) ranging from 0 degrees to 50 degrees C: hoopoe larks (Alaemon alaudipes) and Dunn's larks (Eremalauda dunni) live in hot and dry deserts, whereas skylarks (Alauda arvensis) and woodlarks (Lullula arborea) occur in temperate mesic areas. Mass-adjusted BMR and TEWL were indistinguishable between hoopoe lark and Dunn's lark and between skylark and woodlark. When grouping the data of the two desert larks in one set and the data of the two mesic larks in another, desert larks are shown to have 43% lower BMR levels and 27% lower TEWL values than the mesic species. Their body temperatures (T(b)'s) were 1.1 degrees C lower, and the minimal dry heat transfer coefficients (h) were 26% below values for the mesic larks. When T(a) exceeded T(b), the h of hoopoe larks and Dunn's larks was high and indistinguishable from h at 40 degrees C, in contrast to the prediction that h should be decreased to minimize heat gain through conductance, convection, or radiation from the environment when T(a) exceeds T(b).     

Kidney mass and relative medullary thickness of rodents in relation to habitat, body size, and phylogeny

We tested the hypotheses that relative medullary thickness (RMT) and kidney mass are positively related to habitat aridity in rodents, after controlling for correlations with body mass. Body mass, mass-corrected kidney mass, mass-corrected RMT, mass-corrected maximum urine concentration, and habitat (scored on a semiquantitative scale of 1-4 to indicate increasing aridity) all showed statistically significant phylogenetic signal. Body mass varied significantly among habitats, with the main difference being that aquatic species are larger than those from other habitats. Mass-corrected RMT and urine concentration showed a significant positive correlation (N=38; conventional r=0.649, phylogenetically independent contrasts [IC] r=0.685), thus validating RMT as a comparative index of urine concentrating ability. RMT scaled with body mass to an exponent significantly less than 0 (N=141 species; conventional allometric slope=-0.145 [95% confidence interval (CI)=-0.172, -0.117], IC allometric slope=-0.132 [95% CI=-0.180, -0.083]). Kidney mass scaled to an exponent significantly less than unity (N=104 species; conventional slope=0.809 [95% CI=0.751, 0.868], IC slope=0.773 [95% CI=0.676, 0.871]). Both conventional and phylogenetic analysis indicated that RMT varied among habitats, with rodents from arid areas having the largest values of RMT. A phylogenetic analysis indicated that mass-corrected kidney mass was positively related to habitat aridity.     

Water and energy economy of an omnivorous bird: population differences in the Rufous-collared Sparrow (Zonotrichia capensis)

We investigated the intraspecific variation in basal metabolic rate (BMR) and total evaporative water loss (TEWL) in the omnivorous passerine Zonotrichia capensis from two populations inhabiting regions with different precipitation regimes and aridity indices. Values of TEWL in birds from the semi-arid region were significantly lower than those found in sparrows from the mesic region. TEWL in birds from the semi-arid site was 74% of the expectation based on body mass for passerines from mesic areas and similar to the allometric expectation for passerines from arid environments. In sparrows from the mesic area, TEWL was higher than predicted by their body mass for passerines from arid environments (133%), but very close (97%) to the expectation for passerines from mesic areas. BMR values were 25% lower in sparrows from the semi-arid region. The lower TEWL and BMR of birds from the semi-arid region may be a physiological adjustment that allows them to cope with fewer resources and/or water. We propose that the lower endogenous heat production in birds from the semi-arid environment may decrease their water requirements.     

Environmental correlates of physiological variables in marsupials

We analyzed body temperature (T(b)), basal metabolic rate (BMR), wet thermal conductance (C(wet)), and evaporative water loss (EWL) of marsupials by conventional and phylogenetically corrected regression. Allometric effects were substantial for BMR, C(wet), and EWL but not T(b). There was a strong phylogenetic signal for mass and all physiological traits. A significant phylogenetic signal remained for BMR, C(wet), and EWL even after accounting for the highly significant phylogenetic signal of mass. T(b), BMR, C(wet), and EWL allometric residuals were correlated with some diet, distribution, and climatic variables before and after correction for phylogeny. T(b) residuals were higher for marsupials from arid environments (high T(a) and more variable rainfall). The fossorial marsupial mole had a lower-than-expected T(b) residual. The allometric slope for BMR was 0.72-0.75. Residuals were consistently related to distribution aridity and rainfall variability, with species from arid and variable rainfall habitats having a low BMR, presumably to conserve energy in a low-productivity environment. The nectarivorous honey possum had a higher-than-expected BMR. For C(wet), the allometric slope was 0.55-0.62; residuals were related to diet, with folivores having low and insectivores high C(wet) residuals. The allometric slope for EWL was 0.68-0.73. EWL residuals were consistently correlated with rainfall variability, presumably facilitating maintenance of water balance during dry periods.     

Ecological and evolutionary physiology of desert birds: a progress report

The adaptive significance of mechanisms of energy and water conservation among species of desert rodents, which avoid temperature extremes by remaining within a burrow during the day, is well established. Conventional wisdom holds that arid-zone birds, diurnal organisms that endure the brunt of their environment, occupy these desert climates because of the possession of physiological design features common to all within the class Aves. We review studies that show that desert birds may have evolved specific features to deal with hot desert conditions including: a reduced basal metabolic rate (BMR) and field metabolic rate (FMR), and lower total evaporative water loss (TEWL) and water turnover (WTO).Previous work on the comparative physiology of desert birds relied primarily on information gathered on species from the deserts of the southwestern U.S., which are semi-arid habitats of recent geologic origin. We include data on species from Old World deserts, which are geologically older than those in the New World, and place physiological responses along an aridity axis that includes mesic, semi-arid, arid, and hyperarid environments.The physiological differences between desert and mesic birds that we have identified using the comparative method could arise as a result of acclimation to different environments, of genetic change mediated by selection, or both. We present data on the flexibility of BMR and TEWL in Hoopoe Larks that suggest that phenotypic adjustments in these variables can be substantial. Finally, we suggest that linkages between the physiology of individual organism and its life-history are fundamental to the understanding of life-history evolution.     

Diet, phylogeny, and basal metabolic rate in phyllostomid bats

Aside from the pervasive effects of body mass, much controversy exists as to what factors account for interspecific variation in basal metabolic rates (BMR) of mammals; however, both diet and phylogeny have been strongly implicated. We examined variation in BMR within the New World bat family Phyllostomidae, which shows the largest diversity of food habits among mammalian families, including frugivorous, nectarivorous, insectivorous, carnivorous and blood-eating species. For 27 species, diet was taken from the literature and BMR was either measured on animals captured in Brazil or extracted from the literature. Conventional (nonphylogenetic) analysis of covariance (ANCOVA), with body mass as the covariate, was first used to test the effects of diet on BMR. In this analysis, which assumes that all species evolved simultaneously from a single ancestor (i.e., a "star" phylogeny), diet exerted a strong effect on mass-independent BMR: nectarivorous bats showed higher mass-independent BMR than other bats feeding on fruits, insects or blood. In phylogenetic ANCOVAs via Monte Carlo computer simulation, which assume that species are part of a branching hierarchical phylogeny, no statistically significant effect of diet on BMR was observed. Hence, results of the nonphylogenetic analysis were misleading because the critical values for testing the effect of diet were underestimated. However, in this sample of bats, diet is perfectly confounded with phylogeny, because the four dietary categories represent four separate subclades, which greatly reduces statistical power to detect a diet (= subclade) effect. But even if diet did appear to exert an influence on BMR in this sample of bats, it would not be logically possible to separate this effect from the possibility that the dietary categories differ for some other reason (i.e., another synapomorphy of one or more of the subclades). Examples such as this highlight the importance of considering phylogenetic relationships when designing new comparative studies, as well as when analyzing existing data sets. We also discuss some possible reasons why BMR may not coadapt with diet.     

Environment, migratory tendency, phylogeny and basal metabolic rate in birds

Basal metabolic rate (BMR) represents the minimum maintenance energy requirement of an endotherm and has far-reaching consequences for interactions between animals and their environments. Avian BMR exhibits considerable variation that is independent of body mass. Some long-distance migrants have been found to exhibit particularly high BMR, traditionally interpreted as being related to the energetic demands of long-distance migration. Here we use a global dataset to evaluate differences in BMR between migrants and non-migrants, and to examine the effects of environmental variables. The BMR of migrant species is significantly higher than that of non-migrants. Intriguingly, while the elevated BMR of migrants on their breeding grounds may reflect the metabolic machinery required for long-distance movements, an alternative (and statistically stronger) explanation is their occupation of predominantly cold high-latitude breeding areas. Among several environmental predictors, average annual temperature has the strongest effect on BMR, with a 50% reduction associated with a 20 degrees C gradient. The negative effects of temperature variables on BMR hold separately for migrants and non-migrants and are not due their different climatic associations. BMR in migrants shows a much lower degree of phylogenetic inertia. Our findings indicate that migratory tendency need not necessarily be invoked to explain the higher BMR of migrants. A weaker phylogenetic signal observed in migrants supports the notion of strong phenotypic flexibility in this group which facilitates migration-related BMR adjustments that occur above and beyond environmental conditions. In contrast to the findings of previous analyses of mammalian BMR, primary productivity, aridity or precipitation variability do not appear to be important environmental correlates of avian BMR. The strong effects of temperature-related variables and varying phylogenetic effects reiterate the importance of addressing both broad-scale and individual-scale variation for understanding the determinants of BMR.     

Climatic adaptation and the evolution of basal and maximum rates of metabolism in rodents

Metabolic rate is a key aspect of organismal biology and the identification of selective factors that have led to species differences is a major goal of evolutionary physiology. We tested whether environmental characteristics and/or diet were significant predictors of interspecific variation in rodent metabolic rates. Mass-specific basal metabolic rates (BMR) and maximum metabolic rates (MMR, measured during cold exposure in a He-O2 atmosphere) were compiled from the literature. Maximum (Tmax) and minimum (Tmin) annual mean temperatures, latitude, altitude, and precipitation were obtained from field stations close to the capture sites reported for each population (N = 57). Diet and all continuous-valued traits showed statistically significant phylogenetic signal, with the exception of mass-corrected MMR and altitude. Therefore, results of phylogenetic analyses are emphasized. Body mass was not correlated with absolute latitude, but was positively correlated with precipitation in analyses with phylogenetically independent contrasts. Conventional multiple regressions that included body mass indicated that Tmax (best), Tmin, latitude, and diet were significant additional predictors of BMR. However, phylogenetic analyses indicated that latitude was the only significant predictor of mass-adjusted BMR (positive partial regression coefficient, one-tailed P = 0.0465). Conventional analyses indicated that Tmax, Tmin (best), and altitude explained significant amounts of the variation in mass-adjusted MMR. With body mass and Tmin in the model, no additional variables were significant predictors. Phylogenetic contrasts yielded similar results. Both conventional and phylogenetic analyses indicated a highly significant positive correlation between residual BMR and MMR (as has also been reported for birds), which is consistent with a key assumption of the aerobic capacity model for the evolution of vertebrate energetics (assuming that MMR and exercise-induced maximal oxygen consumption are positively functionally related). Our results support the hypothesis that variation in environmental factors leads to variation in the selective regime for metabolic rates of rodents. However, the causes of a positive association between BMR and latitude remain obscure. Moreover, an important area for future research will be experiments in all taxa are raised under common conditions to allow definitive tests of climatic adaptation in endotherm metabolic rates and to elucidate the extent of adaptive phenotypic plasticity.     

A humped latitudinal phylogenetic diversity pattern of orchid bees (Hymenoptera: Apidae: Euglossini) in western Amazonia: assessing the influence of climate and geologic history

Amazonian rainforests are among the most species‐rich terrestrial habitats on Earth. The aim of this study was to analyze phylogenetic diversity (PD) patterns of orchid bee assemblages along a latitudinal gradient of 15° from northern Peru to central Bolivia and to relate them to climatic factors and geological history. We expanded an existing phylogeny of orchid bees and analyzed the PD of 15 orchid bee assemblages along a latitudinal gradient using mean pair‐wise phylogenetic distance. The resulting pattern was correlated to climatic factors and elevation. We found a hump‐shaped pattern of PD that peaked in central Peru and decreased towards the equatorial and especially towards the southern, subtropical sites. The decrease in PD towards higher latitudes is a common pattern found in many taxa, which in our case correlates with increasing climatic seasonality. However, the decrease towards the equatorial sites is unusual and may be related to a particular historic event: the northern sites with low PD are situated in the area of the former Lake Pebas, which covered western Amazonia until 3 mya. After the lake disappeared orchid bees mainly belonging to two distantly related species groups apparently colonized the region and diversified, which led to the comparatively low observed PD. In contrast, in central Peru, no in situ radiations were detected, hence the assemblages were composed of species from diverse phylogenetic lineages. Additionally, we identified multiple phylogenetically independent radiations of regionally restricted Euglossa species along the latitudinal gradient that, according to a published, dated phylogeny, diversified roughly 3–1 mya. The hump‐shaped latitudinal pattern of PD of the orchid bees of western Amazonia thus appears to have resulted from a preponderance of early divergent lineages in central Peru and of young radiations from distantly related clades colonizing higher latitudes, possibly triggered by historic climate fluctuations and major geological events.     

Using the Past to Predict the Present: Confidence Intervals for Regression Equations in Phylogenetic Comparative Methods

Two phylogenetic comparative methods, independent contrasts and generalized least squares models, can be used to determine the statistical relationship between two or more traits. We show that the two approaches are functionally identical and that either can be used to make statistical inferences about values at internal nodes of a phylogenetic tree (hypothetical ancestors), to estimate relationships between characters, and to predict values for unmeasured species. Regression equations derived from independent contrasts can be placed back onto the original data space, including computation of both confidence intervals and prediction intervals for new observations. Predictions for unmeasured species (including extinct forms) can be made increasingly accurate and precise as the specificity of their placement on a phylogenetic tree increases, which can greatly increase statistical power to detect, for example, deviation of a single species from an allometric prediction. We reexamine published data for basal metabolic rates (BMR) of birds and show that conventional and phylogenetic allometric equations differ significantly. In new results, we show that, as compared with nonpasserines, passerines exhibit a lower rate of evolution in both body mass and mass-corrected BMR; passerines also have significantly smaller body masses than their sister clade. These differences may justify separate, clade-specific allometric equations for prediction of avian basal metabolic rates.     

Patterns of Ant (Hymenoptera: Formicidae) Richness and Relative Abundance along an Aridity Gradient in Western Venezuela

In xeric ecosystems, ant diversity response to aridity varies with rainfall magnitude and gradient extension. At a local scale and with low precipitation regimes, increased aridity leads to a reduction of species richness and an increased relative abundance for some ant species. In order to test this pattern in tropical environments, ant richness and relative abundance variation were evaluated along 35 km of an aridity gradient in the Araya Peninsula, state of Sucre, Venezuela. Three sampling stations comprising five transects each were set up. Pitfall traps and direct collecting from vegetation were assessed per transect. Overall, 52 species, 23 genera, and 7 subfamilies of ants were recorded in the peninsula. The total number of species and genera recorded by both sampling stations and transects decreased linearly with increasing aridity. Total relative abundance was highest in the most arid portion of the peninsula, with Crematogaster rochai (Forel) and Camponotus conspicuus zonatus (Emery) (Hymenoptera: Formicidae) being the numerically dominant species. Spatial and multivariate analyses revealed significant changes in ant composition every 11 km of distance, and showed a decrease of ant diversity with the increase of harsh conditions in the gradient. Here, we discuss how local geographic and topographic features of Araya originate the aridity gradient and so affect the microhabitat conditions for the ant fauna.     

Constraints on trait combinations explain climatic drivers of biodiversity: the importance of trait covariance in community assembly

Trade‐offs maintain diversity and structure communities along environmental gradients. Theory indicates that if covariance among functional traits sets a limit on the number of viable trait combinations in a given environment, then communities with strong multidimensional trait constraints should exhibit low species diversity. We tested this prediction in winter annual plant assemblages along an aridity gradient using multilevel structural equation modelling. Univariate and multivariate functional diversity measures were poorly explained by aridity, and were surprisingly poor predictors of community richness. By contrast, the covariance between maximum height and seed mass strengthened along the aridity gradient, and was strongly associated with richness declines. Community richness had a positive effect on local neighbourhood richness, indicating that climate effects on trait covariance indirectly influence diversity at local scales. We present clear empirical evidence that declines in species richness along gradients of environmental stress can be due to increasing constraints on multidimensional phenotypes.     

Physiological responses in rufous-collared sparrows to thermal acclimation and seasonal acclimatization

A large number of physiological acclimation studies assume that flexibility in a certain trait is both adaptive and functionally important for organisms in their natural environment; however, it is not clear how an organism’s capacity for temperature acclimation translates to the seasonal acclimatization that these organisms must accomplish. To elucidate this relationship, we measured BMR and TEWL rates in both field-acclimatized and laboratory-acclimated adult rufous-collared sparrows (Zonotrichia capensis). Measurements in field-acclimatized birds were taken during the winter and summer seasons; in the laboratory-acclimated birds, we took our measurements following 4 weeks at either 15 or 30°C. Although BMR and TEWL rates did not differ between winter and summer in the field-acclimatized birds, laboratory-acclimated birds exposed to 15°C exhibited both a higher BMR and TEWL rate when compared to the birds acclimated to 30°C and the field-acclimatized birds. Because organ masses seem to be similar between field and cold-acclimated birds whereas BMR is higher in cold-acclimated birds, the variability in BMR cannot be explained completely by adjustments in organ masses. Our findings suggest that, although rufous-collared sparrows can exhibit thermal acclimation of physiological traits, sparrows do not use this capacity to cope with minor to moderate fluctuations in environmental conditions. Our data support the hypothesis that physiological flexibility in energetic traits is a common feature of avian metabolism.     

Repeatability and individual correlates of basal metabolic rate and total evaporative water loss in birds: a case study in European stonechats

Basal metabolic rate (BMR) and total evaporative water loss (TEWL) are thought to have evolved in conjunction with life history traits and are often assumed to be characteristic features of an animal. Physiological traits can show large intraindividual variation at short and long timescales, yet natural selection can only act on a trait if it is a characteristic feature of an individual. The repeatability of a trait, a measure of the portion of variance that is caused by differences among individuals, indicates if it is a characteristic feature of an individual. We measured repeatability of BMR and TEWL of 18 captive European stonechats (Saxicola torquata rubicola) within the winter season. Repeatability was 0.56 for BMR and 0.60 for mass-specific BMR. Age and body mass had a significant effect on variation in BMR. Also after accounting for this variation, BMR remained repeatable. TEWL and mass-specific TEWL showed nonsignificant repeatabilities of 0.11 and 0.12, respectively. We conclude that BMR is a characteristic feature of an individual in our population of European stonechats, whereas TEWL is not. We discuss our results in the context of a review of currently available estimates of repeatability of BMR and TEWL for birds.     

Equivocal evidence for a change in balance between selfing and pollinator-mediated reproduction in annual Brassicaceae growing along a rainfall gradient

Because many plants ensure their reproductive success using some level of self-fertilization, it is predicted that the balance between pollinator-mediated pollen transfer and autonomous selfing will be correlated with habitat harshness and pollinator activity. To study these correlations, we used three annual species that differ in their self-incompatibility and compared their reproductive success in the form of fruit production and seed viability, with and without pollinators, along a rainfall gradient and in relation to pollinator activity. We found that pollinator activity decreased with increasing aridity. Although the reproductive success in the absence of pollinators fitted the species’ level of self-incompatibility, no clear trend was detected between pollinator visits or site harshness and relative reproductive success. Only one of the species, Sinapis alba, showed a consistent increase in autonomous selfing with both increased aridity and decreased pollinator visits, a finding that was supported by an increase in the viability of seeds produced in the absence of pollinators. Nevertheless, unlike our prediction, the absolute number of seeds produced per S. alba flower increased with aridity. Our overall findings suggest that despite decreasing pollinator abundance and diversity with increasing aridity, the reproductive success of annual species, even of those with high self-incompatibility, remains relatively high. Although there may be several mechanisms governing these results, the overall constant seed production despite environmental uncertainties may be part of the mechanism by which plants in arid ecosystems buffer environmental variations.     

Two Mediterranean annuals feature high within-population trait variability and respond differently to a precipitation gradient

Intraspecific trait variability plays an important role in species adaptation to climate change. However, it still remains unclear how plants in semi-arid environments respond to increasing aridity. We investigated the intraspecific trait variability of two common Mediterranean annuals (Geropogon hybridus and Crupina crupinastrum) with similar habitat preferences. They were studied along a steep precipitation gradient in Israel similar to the maximum predicted precipitation changes in the eastern Mediterranean basin (i.e. −30% until 2100). We expected a shift from competitive ability to stress tolerance with decreasing precipitation and tested this expectation by measuring key functional traits (canopy and seed release height, specific leaf area, N- and P-leaf content, seed mass). Further, we evaluated generative bet-hedging strategies by different seed traits. Both species showed different responses along the precipitation gradient. C. crupinastrum exhibited only decreased plant height towards aridity, while G. hybridus showed strong trends of generative adaptation to aridity. Different seed trait indices suggest increased bet-hedging of G. hybridus in arid environments. However, no clear trends along the precipitation gradient were observed in leaf traits (specific leaf area and leaf N-/P-content) in both species. Moreover, variance decomposition revealed that most of the observed trait variation (≫50%) is found within populations. The findings of our study suggest that responses to increased aridity are highly species-specific and local environmental factors may have a stronger effect on intraspecific trait variation than shifts in annual precipitation. We therefore argue that trait-based analyses should focus on precipitation gradients that are comparable to predicted precipitation changes and compare precipitation effects to effects of local environmental factors.     

Molecular phylogenetics of the hummingbird genus Coeligena

Advances in the understanding of biological radiations along tropical mountains depend on the knowledge of phylogenetic relationships among species. Here we present a species-level molecular phylogeny based on a multilocus dataset for the Andean hummingbird genus Coeligena. We compare this phylogeny to previous hypotheses of evolutionary relationships and use it as a framework to understand patterns in the evolution of sexual dichromatism and in the biogeography of speciation within the Andes. Previous phylogenetic hypotheses based mostly on similarities in coloration conflicted with our molecular phylogeny, emphasizing the unreliability of color characters for phylogenetic inference. Two major clades, one monochromatic and the other dichromatic, were found in Coeligena. Closely related species were either allopatric or parapatric on opposite mountain slopes. No sister lineages replaced each other along an elevational gradient. Our results indicate the importance of geographic isolation for speciation in this group and the potential interaction between isolation and sexual selection to promote diversification.     

Basal metabolism is correlated with habitat productivity among populations of degus (Octodon degus)

Several competing hypotheses attempt to explain how environmental conditions affect mass-independent basal metabolic rate (BMR) in mammals. One of the most inclusive is the hypothesis that associates BMR with food habits, including habitat productivity. The effects of food habits have been widely investigated at the interspecific level, and variation between individuals and populations has been largely ignored. Intraspecific analysis of physiological traits has the potential to compensate for many pitfalls associated with interspecific analyses and serve as a useful approach for evaluating hypotheses regarding metabolic adaptation. Here we tested the effects of climatic variables (mean annual rainfall = PP, mean annual temperature = T_A), net primary productivity (NPP) and the de Martonne index (DMi) of aridity on mass-independent BMR among four populations of the caviomorph rodent Octodon degus along a geographic gradient in Chile. BMR was measured on animals maintained in a common garden acclimation set-up, thus kept under the same environment and diet quality for at least 6 months. Mass-independent BMR was significantly different among degu populations showing a large intraspecific spread in metabolic rates. A very large fraction of interpopulational variability in mass-independent BMR was explained by NPP, PP and DMi. Our results were conclusive about the effects of habitat productivity on setting the level of mass-independent BMR at the intraspecific–interpopulational level.     

Seed mass and dormancy of annual plant populations and communities decreases with aridity and rainfall predictability

Several theoretical and empirical studies have examined the influence of environmental conditions on seed traits and germination strategies of annual species. A positive relationship between seed mass and dormancy has been described for annuals occupying climatically unpredictable ecosystems. Larger-seeded species tend to have higher seedling survival rates, while dormancy allows a bet-hedging strategy in unpredictable environments. Until now, these ideas have been addressed primarily for only one or a few focal species, without considering differences among populations and communities. The novelty of the present study lies in the population and community-level approach, where a comprehensive seed trait database including 158 annual species occurring along a gradient of rainfall variability and aridity in Israel was used to ask the following question: Does average seed mass and dormancy of annual populations and communities decrease with increasing aridity and rainfall unpredictability?Soil seed bank samples were collected at the end of the summer drought, before the onset of the rains, from four plant communities. Germination was tested under irrigated conditions during three consecutive germination seasons to determine the overall seed germinability in each soil sample. Seed mass was obtained from newly produced seeds collected at the study sites in late spring. The community level results showed that, in contrast to common theoretical knowledge, seed mass and dormancy of the dominant annual species decreased with increasing aridity and rainfall variability. Accordingly, a negative correlation was found between seed mass and seed germination fractions. The present study demonstrates that an analysis of seed traits along climatic gradients is significantly improved by approaches that target both population and community levels simultaneously. A critical evaluation sheds new light upon the selective pressures that act on seed ecology of annuals along a climatic gradient and facilitates formulation of more mechanistic hypotheses about factors governing critical seed traits.     

Aridity increases below-ground niche breadth in grass communities

Aridity is an important environmental filter in the assembly of plant communities worldwide. The extent to which root traits mediate responses to aridity, and how they are coordinated with leaf traits, remains unclear. Here, we measured variation in root tissue density (RTD), specific root length (SRL), specific leaf area (SLA), and seed size within and among thirty perennial grass communities distributed along an aridity gradient spanning 190–540 mm of climatic water deficit (potential minus actual evapotranspiration). We tested the hypotheses that traits exhibited coordinated variation (1) among species, as well as (2) among communities varying in aridity, and (3) functional diversity within communities declines with increasing aridity, consistent with the “stress-dominance” hypothesis. Across communities, SLA and RTD exhibited a coordinated response to aridity, shifting toward more conservative (lower SLA, higher RTD) functional strategies with increasing aridity. The response of SRL to aridity was more idiosyncratic and was independent of variation in SLA and RTD. Contrary to the stress-dominance hypothesis, the diversity of SRL values within communities increased with aridity, while none of the other traits exhibited significant diversity responses. These results are consistent with other studies that have found SRL to be independent of an SLA–RTD axis of functional variation and suggest that the dynamic nature of soil moisture in arid environments may facilitate a wider array of resource capture strategies associated with variation in SRL.