Developmental time and size-related traits in Drosophila buzzatii along an altitudinal gradient from Argentina

Clinal analysis for fitness-related traits provides a well-known approach to investigate adaptive evolution. Several fitness-related traits (developmental time, thorax length, wing length and wing loading) were measured at two laboratory generations (G7 and G33) of D. buzzatii from an altitudinal gradient from northwestern Argentina, where significant thermal differences persist. Developmental time (DT) was positively correlated with altitude of origin of population. Further, DT was negatively correlated with maximal mean temperature at the site of origin of population, and this thermal variable decreases with altitude. Wing loading tended to be larger in highland than in lowland populations, suggesting that flight performance is subject to stronger selection pressure in highland populations. Developmental time showed a significant increase with laboratory generation number. There was no significant correlation between developmental time and body size across populations along the altitudinal cline of DT. This result illustrates that developmental time and body size do not always evolve in the same direction, even though both traits are often positively and genetically correlated in a well-known tradeoff in Drosophila.     

A field test of the stress‐gradient hypothesis along an aridity gradient

Aims: The stress‐gradient hypothesis (SGH) predicts how plant interactions change along environmental stress gradients. We tested the SGH in an aridity gradient, where support for the hypothesis and the specific shape of its response curve is controversial. Location: Almería, Cáceres and Coimbra, three sites in the Iberian Peninsula that encompass the most arid and wet habitats in the distribution range of a nurse shrub species –Retama sphaerocarpa L. (Boiss) – in Europe. Methods: We analysed the effect of Retama on its understorey plant community and measured the biomass and species richness beneath Retama and in gaps. We estimated the frequency (changes in species richness), importance and intensity of the Retama effects, and derived the severity–interaction relationship pattern, analysing how these interaction indices changed along this aridity gradient. Results and conclusions: The intensity and frequency of facilitation by Retama increased monotonically with increasing environmental severity, and the importance tended to have a similar pattern, overall supporting the SGH. Our data did not support predictions from recent revisions of the SGH, which may not apply to whole plant communities like those studied here or when interactions are highly asymmetrical. Facilitation by Retama influenced community composition and species richness to the point that a significant fraction of species found at the most arid end of the gradient were only able to survive beneath the nurse shrub, whereas some of these species were able to thrive in gaps at more mesic sites, highlighting the ecological relevance of facilitation by nurse species in mediterranean environments, especially in the driest sites.     

Developmental acclimation affects clinal variation in stress resistance traits in Drosophila buzzatii

Patterns of clinal genetic variation in Drosophila are often characterized after rearing at constant temperatures. However, clinal patterns might change after acclimation if populations differ in their plastic response to fluctuating environments. We studied longevity, starvation and heat knock‐down resistance after development at either constant or fluctuating temperatures in nine Drosophila buzzatii populations collected along an altitudinal gradient in Tenerife, Spain. Flies that developed at fluctuating temperatures had higher stress resistance despite experiencing a slightly lower average temperature than those at constant temperatures. Genetic variation along the gradient was found in both stress‐resistance traits. Because Q_ST values greatly exceeded F_ST values, genetic drift could not explain this diversification. In general, differences among populations were larger after rearing at fluctuating temperatures, especially in heat knock‐down, for which clinal patterns disappeared when flies were reared at constant temperatures. This result emphasizes the importance of determining whether populations originating from different environments differ in their plastic responses to stress.     

Hypolithic and soil microbial community assembly along an aridity gradient in the Namib Desert

The Namib Desert is considered the oldest desert in the world and hyperarid for the last 5 million years. However, the environmental buffering provided by quartz and other translucent rocks supports extensive hypolithic microbial communities. In this study, open soil and hypolithic microbial communities have been investigated along an East–West transect characterized by an inverse fog-rainfall gradient. Multivariate analysis showed that structurally different microbial communities occur in soil and in hypolithic zones. Using variation partitioning, we found that hypolithic communities exhibited a fog-related distribution as indicated by the significant East–West clustering. Sodium content was also an important environmental factor affecting the composition of both soil and hypolithic microbial communities. Finally, although null models for patterns in microbial communities were not supported by experimental data, the amount of unexplained variation (68–97 %) suggests that stochastic processes also play a role in the assembly of such communities in the Namib Desert.     

Biotic resistance to invasion along an estuarine gradient

Biotic resistance is the ability of native communities to repel the establishment of invasive species. Predation by native species may confer biotic resistance to communities, but the environmental context under which this form of biotic resistance occurs is not well understood. We evaluated several factors that influence the distribution of invasive Asian mussels (Musculista senhousia) in Mission Bay, a southern California estuary containing an extensive eelgrass (Zostera marina) habitat. Asian mussels exhibit a distinct spatial pattern of invasion, with extremely high densities towards the back of Mission Bay (up to 4,000 m⁻²) in contrast with near-complete absence at sites towards the front of the bay. We established that recruits arrived at sites where adult mussels were absent and found that dense eelgrass does not appear to preclude Asian mussel growth and survival. Mussel survival and growth were high in predator-exclusion plots throughout the bay, but mussel survival was low in the front of the bay when plots were open to predators. Additional experiments revealed that consumption by spiny lobsters (Panulirus interruptus) and a gastropod (Pteropurpura festiva) likely are the primary factors responsible for resistance to Asian mussel invasion. However, biotic resistance was dependent on location within the estuary (for both species) and also on the availability of a hard substratum (for P. festiva). Our findings indicate that biotic resistance in the form of predation may be conferred by higher order predators, but that the strength of resistance may strongly vary across estuarine gradients and depend on the nature of the locally available habitat.     

Life history variation in an annual plant under two opposing environmental constraints along an aridity gradient

Environmental gradients represent an ideal framework for studying adaptive variation in the life history of plant species. However, on very steep gradients, largely contrasting conditions at the two gradient ends often limit the distribution of the same species across the whole range of environmental conditions. Here, we study phenotypic variation in a winter annual crucifer Biscutella didyma persisting along a steep gradient of increasing rainfall in Israel. In particular, we explored whether the life history at the arid end of the gradient indicates adaptations to drought and unpredictable conditions, while adaptations to the highly competitive environment prevail at the mesic Mediterranean end. We examined several morphological and reproductive traits in four natural populations and in populations cultivated in standard common environment. Plants from arid environments were faster in phenological development, more branched in architecture and tended to maximize reproduction, while the Mediterranean plants invested mainly in vertical vegetative growth. Differences between cultivation and field in diaspore production were very large for arid populations as opposed to Mediterranean ones, indicating a larger potential to increase reproduction under favorable conditions. Our overall findings indicate two strongly opposing selective forces at the two extremes of the aridity gradient, which result in contrasting strategies within the studied annual plant species.     

Evolvability of between‐year seed dormancy in populations along an aridity gradient

Under global climate change, adaptation to new conditions is crucial for plant species persistence. This requires the ability to evolve in traits that are correlated with changing climatic variables. We studied between‐year seed dormancy, which correlates with environmental variability, and tested for clinal trends in its evolvability along an aridity gradient in Israel. We conducted a germination experiment under five irrigation levels with two dryland winter annuals (Biscutella didyma, Bromus fasciculatus) from four sites along the gradient. Species differed in means and evolvability of dormancy. Biscutella had high dormancy, which significantly increased with aridity but decreased with higher irrigation. In Bromus, dormancy was low, similar among populations, and only marginally affected by irrigation. Evolvability in Biscutella was high and varied among populations, without a clinal trend along the gradient. Conversely, in Bromus, trait evolvability was low and declined with increasing aridity. We argue that changes in evolvability along climatic gradients depend on the relative intensity of stabilizing selection. This may be high in Bromus and not only depends on environmental stress, but also on variability. Our findings point to the importance of measuring evolvability of climate‐related traits across different natural and artificial environments and for many coexisting species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 924–934.     

Resistance to thermal stress in adult Drosophila buzzatii: acclimation and variation among populations

Survival after heat stress was examined in adult Drosophila buzzatii from seven localities. Males and females were conditioned by a non-lethal high temperature before exposure to a severe thermal stress. Variable times elapsed between conditioning, either as adults or pupae, and exposure to the stress were used, and experimental times to the stress ranged from 2 to 96 hours. Survival after stress varied among populations, and differences generally were consistent across conditioning treatments and across experiments. Resistance to thermal stress was much higher following one conditioning bout 2–4 hours before exposure to a severe stress than when the time elapsed between conditioning and exposure was increased to 24 or more hours. Significantly more adults survived the stress if conditioned 4 days before exposure to the thermal stress, either as adults or as pupae, than if not conditioned. The rank order of resistance roughly followed that predicted from the climatic conditions of the localities of origin.     

Comparative molecular population genetics of the Xdh locus in the cactophilic sibling species Drosophila buzzatii and D. koepferae

The Xdh (rosy) gene is one of the best studied in the Drosophila genus from an evolutionary viewpoint. Here we analyze nucleotide variation in a 1875-bp fragment of the second exon of Xdh in Argentinian populations of the cactophilic D. buzzatii and its sibling D. koepferae. The major electrophoretic alleles of D. buzzatii not only lack diagnostic amino acids in the region studied but also differ on average from each other by four to 13 amino acid changes. Our data also suggest that D. buzzatii populations belonging to different phytogeographic regions are not genetically differentiated, whereas D. koepferae exhibits a significant pattern of population structure. The Xdh region studied is twice as polymorphic in D. buzzatii as in D. koepferae. Differences in historical population size or in recombinational environment between species could account for the differences in the level of nucleotide variation. In both species, the Xdh region exhibits a great number of singletons, which significantly departs from the frequency spectrum expected under neutrality for nonsynonymous sites and also for synonymous sites in D. buzzatii. These excesses of singletons could be the signature of a recent population expansion in D. buzzatii, whereas they may be simply explained as the result of negative selection in D. koepferae.     

Resistance to thermal stress in preadult Drosophila buzzatii: variation among populations and changes in relative resistance across life stages

Resistance to a short term exposure to a high temperature stress was examined in eggs, larvae and pupae of Drosophila buzzfltii from seven localities. Across development, pupae were most resistant, followed by eggs, and then first and third-instar larvae. Variation among populations for resistance to heat stress was significant in all life stages. However, there was much less variation among populations where measured as eggs and pupae than for both first and third instar larvae. Older larvae showed large changes both in viability and developmental time, while exposure of young larvae to heat stress led to a decline in viability without delayed development. Populations that had the shortest developmental time at 25^oC were relatively the most resistant to heat stress as larvae. High relative resistance at one preadult life stage was not necessarily associated with relatively high resistance at another, or with previous measurements of resistance for adults from these populations. Comparison of populations that were more similar in their pattern of change in resistance across development suggested a relationship with the climate of origin. The possibility that developmental variation in the expression of heat shock proteins may cause variation in resistance to thermal stress for different life stages is discussed.     

Altitudinal variation for stress resistance traits and thermal adaptation in adult Drosophila buzzatii from the New World

Multiple stress resistance traits were investigated in the cactophilic fly Drosophila buzzatii. Adults from seven populations derived from North-Western Argentina were compared with respect to traits relevant for thermal stress resistance and for resistance to other forms of environmental stress. The populations were collected along an altitudinal gradient spanning more than 2000 m in height, showing large climatic differences. The results suggest that knock-down resistance to heat stress, desiccation resistance and Hsp70 expression at a relatively severe stressful temperature best reflect thermal adaptation in this species. Furthermore, cold resistance seemed to be of less importance than heat resistance, at least for the adult life stage, in these populations. Clinal variation in thermal resistance traits over short geographical distances suggests relatively strong adaptive differentiation of the populations. This study provides the first evidence for altitudinal differentiation in stress-related traits, and suggests that Hsp70 expression level can be related to altitudinal clines of heat-stress resistance.     

Bottomland hardwood forest species responses to flooding regimes along an urbanization gradient

Urbanization alters stream hydrology, hence flooding frequency and duration in floodplain wetlands. Potential impacts include shifts in species composition and survival, making restoration and selection of wetland species difficult. Cephalanthus occidentalis, Fraxinus pennsylvanica, and Quercus shumardii seedlings were subjected to experimental flooding regimes typical of floodplain forests in rural and urban settings. Treatments included a rural flood regime with three 7-day floods, an Urban-short flood regime with six 4-day floods, and an Urban-long flood regime with six 10-day floods over a growing season. Specific responses, measured by stem length, leaf area, and leaf, stem, and root biomass, varied between species from different wetland indicator classes. C. occidentalis, a wetland obligate, was well adapted to both urban flooding regimes, whereas productivity of F. pennsylvanica, a facultative wetland species, and Q. shumardii, a facultative species, was significantly reduced by the Urban-long treatment. Growth rates also varied over time, indicating the importance of temporal flooding patterns on species productivity. Because urban flooding regimes directly and selectively alter species productivity, proper restoration methods in urbanizing environments should include species selection based on current and potential future hydrologic conditions and use of reference standards from reference sites subjected to similar urban hydrologic regimes.     

Desiccation resistance in four Drosophila species: sex and population effects

Desiccation resistance and body mass were measured in multiple populations of each of four species of Drosophila: two desert endemic species (D. nigrospiracula and D. mojavensis), and two with more widespread distributions (D. melanogaster and D. pseudoobscura). While flies from the desert species were more desiccation tolerant, there was, in certain cases, significant variation in desiccation resistance among populations of the same species. A significant difference in desiccation resistance was observed between the sexes, females were more resistant than males, but this relationship was reversed when taking into account body mass differences between the sexes. The degree of observed within-species variability demonstrates that studies focusing upon differences between species can produce different conclusions if they rely on observations for only single populations of a given species. Our data also suggest the existence of multiple mechanisms for desiccation resistance.     

Summer annual plants exhibit germination responses to an aridity gradient consistent with both delayed germination and phenotypic plasticity

Plant Ecology - Two adaptive strategies to face variable environments have been proposed for annual plants in deserts: delayed and plastic (predictive) germination. All theories and models have...     

Correlated patterns of variation in phenology and seed production in populations of two annual grasses along an aridity gradient

I applied a comparative approach to reveal correlated patterns of variation in phenology and seed production in four populations of two annual grasses Hordeum spontaneum and Avena sterilis, sampled in the same environments distributed along an aridity gradient in Israel. The steep aridity gradient in Israel represents two parallel clines of environmental productivity (annual rainfall) and predictability (variation in amount and timing of annual rainfall) that is likely to induce similar responses in natural plant populations distributed along the gradient, if (1) selection is strong, (2) species share the same ecological niche, and (3) there is genetic variation for ecologically important traits. I found in plants of both species (1) ultimate advance in onset of flowering, and (2) more but smaller seeds, with increasing aridity. The broad sense heritabilities of onset of flowering, seed size and seed yield in both species were very high, moderate and low, respectively. It appears that the observed adaptive complex of traits have evolved in both species in response to this specific array of environments.     

Temperature adaptation of soil bacterial communities along an Antarctic climate gradient: predicting responses to climate warming

Soil microorganisms, the central drivers of terrestrial Antarctic ecosystems, are being confronted with increasing temperatures as parts of the continent experience considerable warming. Here we determined short‐term temperature dependencies of Antarctic soil bacterial community growth rates, using the leucine incorporation technique, in order to predict future changes in temperature sensitivity of resident soil bacterial communities. Soil samples were collected along a climate gradient consisting of locations on the Antarctic Peninsula (Anchorage Island, 67 °34′S, 68 °08′W), Signy Island (60 °43′S, 45 °38′W) and the Falkland Islands (51 °76′S 59 °03′W). At each location, experimental plots were subjected to warming by open top chambers (OTCs) and paired with control plots on vegetated and fell‐field habitats. The bacterial communities were adapted to the mean annual temperature of their environment, as shown by a significant correlation between the mean annual soil temperature and the minimum temperature for bacterial growth (T_min). Every 1 °C rise in soil temperature was estimated to increase T_min by 0.24–0.38 °C. The optimum temperature for bacterial growth varied less and did not have as clear a relationship with soil temperature. Temperature sensitivity, indicated by Q₁₀ values, increased with mean annual soil temperature, suggesting that bacterial communities from colder regions were less temperature sensitive than those from the warmer regions. The OTC warming (generally <1 °C temperature increases) over 3 years had no effects on temperature relationship of the soil bacterial community. We estimate that the predicted temperature increase of 2.6 °C for the Antarctic Peninsula would increase T_min by 0.6–1 °C and Q₁₀ (0–10 °C) by 0.5 units.     

Effects of oxygen on responses to heating in two lizard species sampled along an elevational gradient

Thermal tolerance is an important variable in predictive models about the effects of global climate change on species distributions, yet the physiological mechanisms responsible for reduced performance at high temperatures in air-breathing vertebrates are not clear. We conducted an experiment to examine how oxygen affects three variables exhibited by ectotherms as they heat—gaping threshold, panting threshold, and loss of righting response (the latter indicating the critical thermal maximum)—in two lizard species along an elevational (and therefore environmental oxygen partial pressure) gradient. Oxygen partial pressure did not impact these variables in either species. We also exposed lizards at each elevation to severely hypoxic gas to evaluate their responses to hypoxia. Severely low oxygen partial pressure treatments significantly reduced the gaping threshold, panting threshold, and critical thermal maximum. Further, under these extreme hypoxic conditions, these variables were strongly and positively related to partial pressure of oxygen. In an elevation where both species overlapped, the thermal tolerance of the high elevation species was less affected by hypoxia than that of the low elevation species, suggesting the high elevation species may be adapted to lower oxygen partial pressures. In the high elevation species, female lizards had higher thermal tolerance than males. Our data suggest that oxygen impacts the thermal tolerance of lizards, but only under severely hypoxic conditions, possibly as a result of hypoxia-induced anapyrexia.