Topographic microclimates drive microhabitat associations at the range margin of a butterfly

The habitat associations of individuals underpin the dynamics of species distributions. Broad‐scale gradients in climate can alter habitat associations across species’ geographic ranges, but topographic heterogeneity creates local microclimates which could generate variation in habitat use at finer spatial scales. We examined the selection of microhabitats for egg‐laying by populations of a thermally‐constrained butterfly, the skipper Hesperia comma, across 16 sites with different regional temperatures and topographic microclimates. Using models of thermal microclimate, we examined how the association between eggs and warm bare ground microhabitats varied with ambient temperature, and predicted bare ground associations in 287 existing H. comma populations, to investigate the relative impacts of regional temperatures and topographic microclimates on microhabitat use. Eggs were most strongly associated with bare ground in relatively cool sites, indicating climate‐driven changes in microhabitat use. The majority of temperature variation between study sites was attributable to topographic microclimates rather than regional temperature differences, such that changes in microhabitat associations occurred principally between north‐ and south‐facing slopes within the same region. Predicted microhabitat associations across the UK distribution of H. comma showed that, due to the large temperature differences generated by topography, most of the between‐population variation in microhabitat use occurs locally within 5 km grid squares, with a smaller proportion occurring at a regional level between 5 km squares. Our findings show how microclimatic variation generated by topography alters the habitat associations of populations at fine spatial scales, suggesting that microclimate‐driven changes in habitat suitability could shape species’ distribution dynamics and their responses to environmental change.     

Neither species geographic range size,climatic envelope,nor intraspecific leaf trait variability capture habitat specialization in a hyperdiverse Amazonian forest

Many plant species exhibit strong association with topographic habitats at local scales. However, the historical biogeographic and physiological drivers of habitat specialization are still poorly understood, and there is a need for relatively easy‐to‐measure predictors of species habitat niche breadth. Here, we explore whether species geographic range, climatic envelope, or intraspecific variability in leaf traits is related to the degree of habitat specialization in a hyperdiverse tropical tree community in Amazonian Ecuador. Contrary to our expectations, we find no effect of the size of species geographic ranges, the diversity of climate a species experiences across its range, or intraspecific variability in leaf traits in predicting topographic habitat association in the ~300 most common tropical tree species in a 25‐ha tropical forest plot. In addition, there was no phylogenetic signal to habitat specialization. We conclude that species geographic range size, climatic niche breadth, and intraspecific variability in leaf traits fail to capture the habitat specialization patterns observed in this highly diverse tropical forest.     

Phenotypic plasticity of desiccation resistance in Glossina puparia: are there ecotype constraints on acclimation responses?

Phenotypic plasticity allows organisms to cope with environmental variation and may aid in the evolution of novel traits. However, whether phenotypic plasticity is beneficial, or if acclimation responses might be constrained to particular ecotypes is generally poorly explored. Here we test the beneficial acclimation hypothesis (BAH) and its alternatives for desiccation resistance to atmospheric moisture in mesic‐ and xeric‐adapted Glossina species. Highly significant interactions among acclimation and test humidity were detected for water loss rates indicative of significant phenotypic plasticity. Ordered‐factor anova was unable to reject predictions of the ‘drier is better’ acclimation hypothesis in xeric Glossina morsitans and mesic G. austeni. Evidence for the ‘deleterious acclimation hypothesis’ was found for mesic G. palpalis as expected from the moist habitats it typically occupies. By contrast, support for the ‘optimal acclimation hypothesis’ was found in xeric G. pallidipes. Little support for BAH was obtained in the present study, although other hypotheses, which might enhance fitness within the environments these species are typically exposed to, were supported. However, acclimation responses were not necessarily constrained to xeric/mesic ecotypes which might be expected if adaptation to a particular environment arose as a trade‐off between plastic responses and living in a particular habitat. These results highlight the complexity of acclimation responses and suggest an important role for phenotypic plasticity in moderating environmental effects on evolutionary fitness in Glossina.     

Response of nitrogen cycling to simulated climate change: differential responses along a subalpine ecotone

In situ nitrogen (N) transformations and N availability were examined over a four‐year period in two soil microclimates (xeric and mesic) under a climate‐warming treatment in a subalpine meadow/sagebrush scrub ecotone. Experimental plots that spanned the two soil microclimates were exposed to an in situ infrared (IR) climate change manipulation at the Rocky Mountain Biological Laboratory, near Crested Butte, Colorado. Although the two microclimates did not differ significantly in their rates of N transformations in the absence of heating, they differed significantly in their response to increased IR. Under a simulated warming in the sagebrush‐dominated xeric microclimate, gross N mineralization rates doubled and immobilization rates increased by up to 60% over the first 2 years of the study but declined to predisturbance rates by the fourth year. This temporal pattern of gross mineralization rates correlated with a decline in SOM. Concurrently, rates of net mineralization rates in the heated plots were 60% higher than the controls after the first year. There were no differences in gross or net nitrification rates with heating in the xeric soils. In contrast to the xeric microclimate, there were no significant effects of heating on any N transformation rates in the mesic microclimate. The differing responses in N cycling rates of the two microclimate to the increased IR is most certainly the result of differences in initial soil moisture conditions and vegetation type and cover.     

Revisiting water loss in insects: a large scale view

Desiccation resistance in insects has long been thought to covary with environmental water availability, and to involve changes in both cuticular and respiratory transpiration. Here, we adopt a large-scale approach to address both issues. Water loss rate and precipitation are positively related at global scales. A significant proportion (68%) of the interspecific variation in water loss rate is explained at the genus level or above. The relationship between metabolic rate and water loss rate differs substantially between mesic and xeric species. While these variables covary as a consequence of their independent covariation with body mass in mesic species, this is not the case in xeric species. In the latter, there is a strong relationship between the residuals of the water loss rate-body mass and metabolic rate-body mass relationships, and water loss rate is much reduced. Moreover, because metabolic rate does not differ significantly between xeric and mesic species of a similar size, respiratory transpiration constitutes a greater proportion of total water loss in xeric than in mesic species of a similar size. This implies that respiratory transpiration and the extent to which it can be modified must be of considerable importance in xeric insect species, although finer scale studies suggest otherwise.     

Relation between Water Balance and Climatic Variables Associated with the Geographical Distribution of Anurans

Amphibian species richness increases toward the equator, particularly in humid tropical forests. This relation between amphibian species richness and environmental water availability has been proposed to be a consequence of their high rates of evaporative water loss. In this way, traits that estimate water balance are expected to covary with climate and constrain a species’ geographic distribution. Furthermore, we predicted that coexisting species of anurans would have traits that are adapted to local hydric conditions. We compared the traits that describe water balance in 17 species of anurans that occur in the mesic Atlantic Forest and xeric Cerrado (savannah) habitats of Brazil. We predicted that species found in the warmer and dryer areas would show a lower sensitivity of locomotor performance to dehydration (SLPD), increased resistance to evaporative water loss (REWL) and higher rates of water uptake (RWU) than species restricted to the more mesic areas. We estimated the allometric relations between the hydric traits and body mass using phylogenetic generalized least squares. These regressions showed that REWL scaled negatively with body mass, whereas RWU scaled positively with body mass. Additionally, species inhabiting areas characterized by higher and more seasonally uniform temperatures, and lower and more seasonally concentrated precipitation, such as the Cerrado, had higher RWU and SLPD than species with geographical distributions more restricted to mesic environments, such as the Atlantic Forest. These results support the hypothesis that the interspecific variation of physiological traits shows an adaptation pattern to abiotic environmental traits.     

Patterns of functional clonal traits and clonal growth modes in contrasting grasslands in the central Apennines,Italy

Abstract. Aim: Patterns of plant functional traits related to clonality (clonal growth modes; CGM) in plant communities were studied and hypotheses on the importance of the selected traits in plant communities supported by soils differing in moisture and nutrient status were tested. Material and Methods: Selected plant functional traits, such as the position of the mother‐daughter plants connections, length of spacers, frequency of multiplication, persistency of ramets connections, presence of storage organs and bud protection were studied in two contrasting plant communities (xeric and mesic abandoned pastures) typical of central Apennines, Italy. Results and Discussion: Clonality was shown to be of great importance in both mesic and xeric grasslands. The major differences between the two communities were due to the dominant CGMs: turf graminoids (having effective protection of growth meristems in dense tussocks) dominated xeric grasslands, while rhizomatous graminoids (typical of competitive resource‐rich environments) dominated mesic grasslands. Below‐ground CGOs (clonal growth organs), shorter spacers, higher multiplication potential, permanent ramet connection, large bud bank and increased importance of bud protection were found to be of importance in water stressed xeric grassland. Contrary to our expectations, the mesic (less stressed) grasslands have the higher number of clonal plants possessing storage organs.     

Patterns and causes of morphological population differentiation in the Tenerife skink, Chalcides viridanus

Recent studies indicate that differential selection between mesic and xeric habitats is the most plausible explanation of within-island geographic variation in the Gran Ganarian skink, Chalcides sexlineatus. The island of Tenerife shows mesic/xeric heterogeneity similar to that on Gran Canaria, so we tested the prediction of parallel geographic patterns of morphological population differentiation in the Tenerife skink, Chalcides viridanus. Geographic variation was found to be complex; patterns of differentiation in body dimensions and scalation show evidence of both mosaic and latitudinal facets. Using randomization and regression methods, significant relationships were found between a hypothesized xeric/mesic model of population differentiation and geographic variation in female body dimensions, as well as with substantial proportions of the geographic variation in male and female scalation and, to a lesser extent, body dimensions. Matrix comparisons based on the entire morphological distance matrices show that the patterns of variation are not concordant with an alternative hypothesis of historical secondary contact, or with an isolation-by-distance model. It is argued that a substantial component of the geographic variation in body dimensions and scalation of C. viridanus is due to differential selection along an aridity gradient.     

Phenotypic plasticity in mesic populations of Pinus pinaster improves resistance to xylem embolism (P50) under severe drought

The objectives of the study were to assess the phenotypic variation in the vulnerability to water stress-induced cavitation (estimated by P₅₀, or the xylem water potential which causes a 50% loss of conductivity) and the trade-offs between P₅₀ and related hydraulic traits, i.e., stem specific conductivity (K _s), slope of the vulnerability curve (slope), wood density and branch size. Variability was examined for six Pinus pinaster populations covering the latitudinal range of the species and plasticity was tested through two provenance-progeny trial sites (xeric/mesic). As expected, the overall values of P₅₀, K _s and branch size decreased in the xeric site. Variation in P₅₀ and K _s among populations was mainly the result of phenotypic plasticity, while wood density was genetically controlled and not affected by the environment. Stress conditions in the xeric site promoted a convergence in P₅₀ and K _s as a result of the high phenotypic plasticity of the populations from mesic origins. In the mesic site, the ranking of populations for cavitation resistance and hydraulic capacity was consistent with the geographic location of the seed source. Higher resistance to cavitation was related to lower K _s, branch size and slope, mainly at the population level, but also as a general trend across individuals. In a warmer and drier climate, there could be a potential selection of Pinus pinaster populations from mesic origins, which showed a great responsiveness and adjustment to drought conditions (similar or higher P₅₀ than the populations from dry origins), in addition to a high wood density and growth.     

Forecasting climate change impacts to plant community composition in the Sonoran Desert region

Hotter and drier conditions projected for the southwestern United States can have a large impact on the abundance and composition of long‐lived desert plant species. We used long‐term vegetation monitoring results from 39 large plots across four protected sites in the Sonoran Desert region to determine how plant species have responded to past climate variability. This cross‐site analysis identified the plant species and functional types susceptible to climate change, the magnitude of their responses, and potential climate thresholds. In the relatively mesic mesquite savanna communities, perennial grasses declined with a decrease in annual precipitation, cacti increased, and there was a reversal of the Prosopis velutina expansion experienced in the 20th century in response to increasing mean annual temperature (MAT). In the more xeric Arizona Upland communities, the dominant leguminous tree, Cercidium microphyllum, declined on hillslopes, and the shrub Fouquieria splendens decreased, especially on south‐ and west‐facing slopes in response to increasing MAT. In the most xeric shrublands, the codominant species Larrea tridentata and its hemiparasite Krameria grayi decreased with a decrease in cool season precipitation and increased aridity, respectively. This regional‐scale assessment of plant species response to recent climate variability is critical for forecasting future shifts in plant community composition, structure, and productivity.     

Variation in vegetation and seed bank in a Chilean semi‐arid community affected by ENSO 1997

Abstract. We followed plant cover and soil seed density of shrubs and herbs in two markedly contrasting years with regard to annual rainfall in two opposite‐facing slopes (mesic vs xeric) and an intervening, relatively wet, ravine of a typical creek in semi‐arid Chile. During the ENSO year 1997 cover of ephemerals increased in all three sites; 43% vs 8% on the xeric slope 75% vs 26% on the mesic slope and 57% vs 32% in the ravine. The number of species was almost three times higher on the xeric slope (27 vs 10 species), increased by 47% on the mesic slope (28 vs 19) and by 14% in the ravine (24 vs 21). Cover of shrubs plus perennial grasses increased from 52% in 1996 to 59% in 1997 on the xeric slope, but no significant cover changes were found on the mesic slope or the ravine. On the xeric slope peak density of seeds was 4500/m² in 1996, while during the ENSO year it was 24000/m². On the mesic slope equivalent values were 3000 and 17000 seeds/m² while in the ravine figures were 8500 and 27000 seeds/m². The strong responses of ephemerals and seed bank to the ENSO‐driven wet year of 1997 demonstrate the importance of this phenomenon in replenishing, recovering and probably maintaining ephemeral vegetation in this region.     

Tectonics,topography, and mammalian diversity

Terrestrial vertebrates show striking changes in species richness across topographic gradients. For mammals, nearly twice as many species per unit area occur in topographically complex regions as in adjacent lowlands. The geological context of this pervasive biogeographic pattern suggests that tectonic processes have a first‐order impact on regional diversity. I evaluate ecological, evolutionary, and historical influences of tectonics and topography on the regional diversity of terrestrial mammals, focusing on the hypothesis that diversification rates are higher in active versus passive tectonic settings. Ten predictions follow from this hypothesis. 1) The timing of peaks in speciation should be congruent with the timescale for tectonic episodes. 2) The rates of speciation and genetic differentiation of populations should be greater for species inhabiting topographically complex regions than spatially continuous landscapes. 3) If topographic complexity per se promotes diversification, then a cluster of young divergences should occur for montane species compared to lowland relatives. 4) Endemism in tectonically active regions should reflect origination within the region rather than range reduction from larger areas. 5) Extinction rates should differ for lineages in tectonically active regions compared to adjacent lowlands. 6) The relationship between local and regional species richness should differ between topographic settings because of higher beta diversity in topographically complex regions. 7) Species originating in topographically complex regions should colonize adjacent lowlands more often than the reverse pattern. 8) North‐south mountain ranges should have higher regional species richness than east‐west mountain ranges. 9) Areas with multiple mountain ranges should have higher regional species richness than comparable areas with single mountain ranges. 10) Global climate changes should affect diversification in tectonically active regions. Research addressing these topics places elevational diversity gradients into a geohistorical context and integrates data from modern biotas and the fossil record.     

Galling arthropod diversity in adjacent swamp forests and restinga vegetation in Rio Grande do Sul, Brazil

Galling arthropods create plant structures inside which they find shelter. Factors acting on galler diversity are still being discussed, with this fauna considered more diverse in xeric than mesic environments (higrothermic stress hypothesis, HSH), and also in more plant diverse sites. Here we compare galler abundance (N), equitability (E), species richness (S) and composition between adjacent restinga (xeric) and swamp forests (mesic) in Parque Estadual de Itapeva (29°21' S, 49°45' W), Rio Grande do Sul, southern Brazil. Five trails, two in swamp forest and three in restingas, were sampled four times each (January/December 2005). After an effort of 60h/person, 621 galled plant individuals belonging to 104 gall morphotypes were recorded. This suggests a high galler diversity for the Park, comparable to the richest places known. No differences were found for N, E or S between restingas and swamp forests. However, faunal composition differs significantly between the vegetation types. The dominant (most abundant) species are different in either vegetation type, and are rare or absent on the other vegetation type. Such species composition analysis is still largely ignored for gallers, and stresses the fact that the HSH cannot explain this pattern, since the latter is based on preferences by the ovipositing galler for xeric sites instead of mesic ones. The two habitats differ in microclimate, but species richness, as would be predicted by the HSH, does not differ. This small scale pattern can perhaps be attributed to biogeographic processes on larger scales, as suggested by the resource synchronisation hypothesis.     

Gene flow across a climatic barrier between hybridizing avian species, California and Gambel's quail (Callipepla californica and C. gambelii)

Allopatric species commonly interbreed in a restricted margin between their ranges. The particular factors that permit interbreeding between species determine the extent of hybridization and its significance for evolution and conservation. Using California quail and Gambel's quail (Callipepla californica and C. gambelii) that naturally hybridize in a narrow region between relatively mesic and xeric environments, I assessed the exchange of genetic and phenotypic traits in relation to vegetative and climatic features (temperature and precipitation) that characterize the area of range overlap, and I examined genetic and phenotypic traits within the hybrid zone over a five-year period in relation to variation in precipitation. Using microsatellite markers, this study reveals that genetic, plumage, and morphometric traits are tightly associated with vegetation, rainfall, and temperature profiles through the abrupt transition from one parental species to the other across the hybrid zone. Results show that the hybrid zone has remained clinal, stationary, and bounded over the five-year study period. There was no evidence of introgression outside the narrow hybrid zone. Interannual climatic fluctuations are associated with internal hybrid zone dynamics but did not alter the shape and position of the zone. A transect through the hybrid zone revealed rapid and episodic genetic mixing within the zone. Possible long-term consequences of this restricted hybridization for the evolution of the two parental species are discussed in the light of changing environments.     

Abiotic determinants of the fynbos/succulent karoo boundary,South Africa

This study investigates the influence of texture, soil moisture and nutrient status on the growth and survival of seedlings of two typical fynbos (Leucadendron pubescens and Passerina vulgaris) and succulent karoo (Ruschia spp.) species, which grow in the boundary zone between these two vegetation types. Seedlings of each species were grown in shalederived and sandstone‐derived soils and under xeric and mesic regimes. Under the xeric regime, the shale‐derived and sandstone‐derived soils represented fine and coarse‐textured soils, respectively. Under the mesic regime, the same soils represented nutrient‐rich and nutrient‐poor soils, respectively. The seedlings of both fynbos species died rapidly under the xeric regime, irrespective of soil type. In contrast, the succulent karoo seedlings survived for over 77 days without water. Under mesic conditions, the fynbos seedlings grew faster than the succulent karoo seedlings, irrespective of soil type. Fynbos seedlings appear to be directly limited by the environment (moisture and salinity), whereas succulent karoo seedlings may be limited by interactions with other plants.     

Native and exotic plant cover vary inversely along a climate gradient 11 years following stand‐replacing wildfire in a dry coniferous forest,Oregon, USA

Community re‐assembly following future disturbances will often occur under warmer and more moisture‐limited conditions than when current communities assembled. Because the establishment stage is regularly the most sensitive to climate and competition, the trajectory of recovery from disturbance in a changing environment is uncertain, but has important consequences for future ecosystem functioning. To better understand how ongoing warming and rising moisture limitation may affect recovery, we studied native and exotic plant composition 11 years following complete stand‐replacing wildfire in a dry coniferous forest spanning a large gradient in climatic moisture deficit (CMD) from warm and dry low elevation sites to relatively cool and moist higher elevations sites. We then projected future precipitation, temperature and CMD at our study locations for four scenarios selected to encompass a broad range of possible future conditions for the region. Native perennials dominated relatively cool and moist sites 11 years after wildfire, but were very sparse at the warmest and driest (high CMD) sites, particularly when combined with high topographic sun exposure. In contrast, exotic species (primarily annual grasses) were dominant or co‐dominant at the warmest and driest sites, especially with high topographic sun exposure. All future scenarios projected increasing temperature and CMD in coming decades (e.g., from 4.5% to 29.5% higher CMD by the 2080's compared to the 1971–2000 average), even in scenarios where growing season (May‐September) precipitation increased. These results suggest increasing temperatures and moisture limitation could facilitate longer term (over a decade) transitions toward exotic‐dominated communities after severe wildfire when a suitable exotic seed source is present.     

Ecological traits influence the phylogenetic structure of bird species co‐occurrences worldwide

The extent to which species’ ecological and phylogenetic relatedness shape their co‐occurrence patterns at large spatial scales remains poorly understood. By quantifying phylogenetic assemblage structure within geographic ranges of >8000 bird species, we show that global co‐occurrence patterns are linked – after accounting for regional effects – to key ecological traits reflecting diet, mobility, body size and climatic preference. We found that co‐occurrences of carnivorous, migratory and cold‐climate species are phylogenetically clustered, whereas nectarivores, herbivores, frugivores and invertebrate eaters tend to be more phylogenetically overdispersed. Preference for open or forested habitats appeared to be independent from the level of phylogenetic clustering. Our results advocate for an extension of the tropical niche conservatism hypothesis to incorporate ecological and life‐history traits beyond the climatic niche. They further offer a novel species‐oriented perspective on how biogeographic and evolutionary legacies interact with ecological traits to shape global patterns of species coexistence in birds.     

Habitat associations of species show consistent but weak responses to climate

Different vegetation types can generate variation in microclimates at local scales, potentially buffering species from adverse climates. To determine if species could respond to such microclimates under climatic warming, we evaluated whether ectothermic species (butterflies) can exploit favourable microclimates and alter their use of different habitats in response to year-to-year variation in climate. In both relatively cold (Britain) and warm (Catalonia) regions of their geographical ranges, most species shifted into cooler, closed habitats (e.g. woodland) in hot years, and into warmer, open habitats (e.g. grassland) in cooler years. Additionally, three-quarters of species occurred in closed habitats more frequently in the warm region than in the cool region. Thus, species shift their local distributions and alter their habitat associations to exploit favourable microclimates, although the magnitude of the shift (approx. 1.3% of individuals from open to shade, per degree Celsius) is unlikely to buffer species from impacts of regional climate warming.     

Range size: disentangling current traits and phylogenetic and biogeographic factors

The range size of a species can be determined by its current traits and by phylogenetic and biogeographic factors. However, only rarely have these factors been studied in combination. We use data on the geographic range sizes of all 26 Sylvia warblers to explicitly test whether range size was determined by current species-specific traits (e.g., body size, dispersal ability), phylogenetic factors (e.g., age of the lineage), or environmental, biogeographic factors (e.g., latitudinal position of the range). The results demonstrated that current traits and phylogenetic and biogeographic factors were interrelated. While a number of factors were significant in simple regression analyses, only one factor determined range size in the multiple regression analyses--dispersal ability. Species with better dispersal ability had larger ranges than species with poorer dispersal ability. Apparent increases of range size with latitude or with the age of the species resulted from correlations with dispersal ability. While the most significant factor that influences the range size of a group of species might differ from one group to the next, these results demonstrate that studies that focus only on a single, for example, phylogenetic, factor might yield misleading results.     

The effect of range changes on the functional turnover,structure and diversity of bird assemblages under future climate scenarios

Animal assemblages fulfill a critical set of ecological functions for ecosystems that may be altered substantially as climate change‐induced distribution changes lead to community disaggregation and reassembly. We combine species and community perspectives to assess the consequences of projected geographic range changes for the diverse functional attributes of avian assemblages worldwide. Assemblage functional structure is projected to change highly unevenly across space. These differences arise from both changes in the number of species and changes in species’ relative local functional redundancy or distinctness. They sometimes result in substantial losses of functional diversity that could have severe consequences for ecosystem health. Range expansions may counter functional losses in high‐latitude regions, but offer little compensation in many tropical and subtropical biomes. Future management of local community function and ecosystem services thus relies on understanding the global dynamics of species distributions and multiscale approaches that include the biogeographic context of species traits.