Virulence‐driven trade‐offs in disease transmission: A meta‐analysis*

The virulence–transmission trade‐off hypothesis proposed more than 30 years ago is the cornerstone in the study of host–parasite co‐evolution. This hypothesis rests on the premise that virulence is an unavoidable and increasing cost because the parasite uses host resources to replicate. This cost associated with replication ultimately results in a deceleration in transmission rate because increasing within‐host replication increases host mortality. Empirical tests of predictions of the hypothesis have found mixed support, which cast doubt about its overall generalizability. To quantitatively address this issue, we conducted a meta‐analysis of 29 empirical studies, after reviewing over 6000 published papers, addressing the four core relationships between (1) virulence and recovery rate, (2) within‐host replication rate and virulence, (3) within‐host replication and transmission rate, and (4) virulence and transmission rate. We found strong support for an increasing relationship between replication and virulence, and replication and transmission. Yet, it is still uncertain if these relationships generally decelerate due to high within‐study variability. There was insufficient data to quantitatively test the other two core relationships predicted by the theory. Overall, the results suggest that the current empirical evidence provides partial support for the trade‐off hypothesis, but more work remains to be done.     

Global change synergies and trade‐offs between renewable energy and biodiversity

Reliance on fossil fuels is causing unprecedented climate change and is accelerating environmental degradation and global biodiversity loss. Together, climate change and biodiversity loss, if not averted urgently, may inflict severe damage on ecosystem processes, functions and services that support the welfare of modern societies. Increasing renewable energy deployment and expanding the current protected area network represent key solutions to these challenges, but conflicts may arise over the use of limited land for energy production as opposed to biodiversity conservation. Here, we compare recently identified core areas for the expansion of the global protected area network with the renewable energy potential available from land‐based solar photovoltaic, wind energy and bioenergy (in the form of Miscanthus × giganteus). We show that these energy sources have very different biodiversity impacts and net energy contributions. The extent of risks and opportunities deriving from renewable energy development is highly dependent on the type of renewable source harvested, the restrictions imposed on energy harvest and the region considered, with Central America appearing at particularly high potential risk from renewable energy expansion. Without restrictions on power generation due to factors such as production and transport costs, we show that bioenergy production is a major potential threat to biodiversity, while the potential impact of wind and solar appears smaller than that of bioenergy. However, these differences become reduced when energy potential is restricted by external factors including local energy demand. Overall, we found that areas of opportunity for developing solar and wind energy with little harm to biodiversity could exist in several regions of the world, with the magnitude of potential impact being particularly dependent on restrictions imposed by local energy demand. The evidence provided here helps guide sustainable development of renewable energy and contributes to the targeting of global efforts in climate mitigation and biodiversity conservation.     

Rapid thermal adaptation in a marine diatom reveals constraints and trade‐offs

Rapid evolution in response to environmental change will likely be a driving force determining the distribution of species across the biosphere in coming decades. This is especially true of microorganisms, many of which may evolve in step with warming, including phytoplankton, the diverse photosynthetic microbes forming the foundation of most aquatic food webs. Here we tested the capacity of a globally important, model marine diatom Thalassiosira pseudonana, for rapid evolution in response to temperature. Selection at 16 and 31°C for 350 generations led to significant divergence in several temperature response traits, demonstrating local adaptation and the existence of trade‐offs associated with adaptation to different temperatures. In contrast, competitive ability for nitrogen (commonly limiting in marine systems), measured after 450 generations of temperature selection, did not diverge in a systematic way between temperatures. This study shows how rapid thermal adaptation affects key temperature and nutrient traits and, thus, a population's long‐term physiological, ecological, and biogeographic response to climate change.     

Phenotypic constraints and community structure: Linking trade‐offs within and among species

Trade‐offs are central to many topics in biology, from the evolution of life histories to ecological mechanisms of species coexistence. Trade‐offs observed among species may reflect pervasive constraints on phenotypes that are achievable given biophysical and resource limitations. If so, then among‐species trade‐offs should be consistent with trade‐offs within species. Alternatively, trait variation among co‐occurring species may reflect historical contingencies during community assembly rather than within‐species constraints. Here, we test whether a key trade‐off between relative growth rate (RGR) and water‐use efficiency (WUE) among Sonoran Desert winter annual plants is apparent within four species representing different strategies in the system. We grew progeny of maternal families from multiple populations in a greenhouse common garden. One species, Pectocarya recurvata, displayed the expected RGR–WUE trade‐off among families within populations. For other species, although RGR and WUE often varied clinally among populations, among‐family variation within populations was lacking, implicating a role for past selection on these traits. Our results suggest that a combination of limited genetic variation in single traits and negative trait correlations could pose constraints on the evolution of a high‐RGR and high‐WUE phenotype within species, providing a microevolutionary explanation for phenotypes that influence community‐level patterns of abundance and coexistence.     

Selection and constraints on offspring size‐number trade‐offs in sand lizards (Lacerta agilis)

The trade‐off between offspring size and number is a central component of life‐history theory, postulating that larger investment into offspring size inevitably decreases offspring number. This trade‐off is generally discussed in terms of genetic, physiological or morphological constraints; however, as among‐individual differences can mask individual trade‐offs, the underlying mechanisms may be difficult to reveal. In this study, we use multivariate analyses to investigate whether there is a trade‐off between offspring size and number in a population of sand lizards by separating among‐ and within‐individual patterns using a 15‐year data set collected in the wild. We also explore the ecological and evolutionary causes and consequences of this trade‐off by investigating how a female's resource (condition)‐ vs. age‐related size (snout‐vent length) influences her investment into offspring size vs. number (OSN), whether these traits are heritable and under selection and whether the OSN trade‐off has a genetic component. We found a negative correlation between offspring size and number within individual females and physical constraints (size of body cavity) appear to limit the number of eggs that a female can produce. This suggests that the OSN trade‐off occurs due to resource constraints as a female continues to grow throughout life and, thus, produces larger clutches. In contrast to the assumptions of classic OSN theory, we did not detect selection on offspring size; however, there was directional selection for larger clutch sizes. The repeatabilities of both offspring size and number were low and we did not detect any additive genetic variance in either trait. This could be due to strong selection (past or current) on these life‐history traits, or to insufficient statistical power to detect significant additive genetic effects. Overall, the findings of this study are an important illustration of how analyses of within‐individual patterns can reveal trade‐offs and their underlying causes, with potential evolutionary and ecological consequences that are otherwise hidden by among‐individual variation.     

Effects of organ removal on trade‐offs between sexual and clonal reproduction in the stoloniferous herb Duchesnea indica

Resource allocation patterns and trade‐off between sexual and clonal reproduction in clonal plants have been extensively studied, but little is known about effects of organ removal on the trade‐offs in clonal plants. To examine the effects, we conducted an experiment with the stoloniferous herb Duchesnea indica in which we removed plant organs like roots, flowers, or fruits. Removing roots significantly increased number of ramets and biomass allocation to stolons, but decreased number of fruits. Removing flowers or fruits greatly increased number of ramets and biomass allocation to stolons and roots, but decreased spacer length, number of fruits, and fruits set. Onset and median date of flowering phenology of D. indica shifted after flowers, fruits, or roots were removed. These results may indicate that removing organs can affect trade‐off between sexual and clonal reproduction of D. indica.     

Local above‐ground persistence of vascular plants: Life‐history trade‐offs and environmental constraints

Questions: 1. Which plant traits and habitat characteristics best explain local above‐ground persistence of vascular plant species and 2. Is there a trade‐off between local above‐ground persistence and the ability for seed dispersal and below‐ground persistence in the soil seed bank? Locations: 845 long‐term permanent plots in terrestrial habitats across the Netherlands. Methods: We analysed the local above‐ground persistence of vascular plants in permanent plots (monitored once a year for ca. 16 year) with respect to functional traits and habitat preferences using survival statistics (Kaplan‐Meier analysis and Cox’ regression). These methods account for censored data and are rarely used in vegetation ecology. Results: Local above‐ground persistence is determined by both functional traits (especially the ability to form long‐lived clonal connections) and habitat preferences (especially nutrient requirements). Above‐ground persistence is negatively related to the ability for dispersal by wind and to the ability to accumulate a long‐term persistent soil seed bank (‘dispersal through time’) and is positively related to the ability for dispersal by water. Conclusions: Most species have a half‐life expectation over 15 years, which may contribute to time lags after changes in habitat quality or ‐configuration (‘extinction debt’). There is evidence for a trade‐off relationship between local above‐ground persistence and below‐ground seed persistence, while the relationship with dispersal in space is vector specific. The rate of species turnover increases with productivity.     

Give ‘til it hurts: trade‐offs between immunity and male reproductive effort in the decorated cricket,Gryllodes sigillatus

Trade‐offs between life‐history variables can be manifested at either the phenotypic or genetic level, with vastly different evolutionary consequences. Here, we examined whether male decorated crickets (Gryllodes sigillatus) from eight inbred lines and the outbred founder population from which they were derived, trade‐off immune effort [lytic activity, phenoloxidase (PO) activity or encapsulation] to produce spermatophylaxes: costly nuptial food gifts essential for successful sperm transfer. Canonical correlation analysis of the outbred population revealed a trade‐off between spermatophylax mass and lytic activity. Analysis of our inbred lines, however, revealed that although PO activity, encapsulation, body mass, spermatophylax mass and ampulla (sperm capsule) mass were all highly heritable, lytic activity was not, and there was, therefore, no negative genetic correlation between lytic activity and spermatophylax mass. Thus, males showed a phenotypic but not a genetic trade‐off between spermatophylax mass and lytic activity, suggesting that this trade‐off is mediated largely by environmental factors.     

Evolutionary constraint on low elevation range expansion: Defense‐abiotic stress‐tolerance trade‐off in crosses of the ecological model Boechera stricta

Most transplant experiments across species geographic range boundaries indicate that adaptation to stressful environments outside the range is often constrained. However, the mechanisms of these constraints remain poorly understood. We used extended generation crosses from diverged high and low elevation populations. In experiments across low elevation range boundaries, there was selection on the parental lines for abiotic stress‐tolerance and resistance to herbivores. However, in support of a defense‐tolerance trade‐off, extended generation crosses showed nonindependent segregation of these traits in the laboratory across a drought‐stress gradient and in the field across the low elevation range boundary. Genotypic variation in a marker from a region of the genome containing a candidate gene (MYC2) was associated with change in the genetic trade‐off. Thus, using crosses and forward genetics, we found experimental genetic and molecular evidence for a pleiotropic trade‐off that could constrain the evolution of range expansion.     

Co‐occurrence patterns of trees along macro‐climatic gradients and their potential influence on the present and future distribution of Fagus sylvatica L.

Aim During recent and future climate change, shifts in large‐scale species ranges are expected due to the hypothesized major role of climatic factors in regulating species distributions. The stress‐gradient hypothesis suggests that biotic interactions may act as major constraints on species distributions under more favourable growing conditions, while climatic constraints may dominate under unfavourable conditions. We tested this hypothesis for one focal tree species having three major competitors using broad‐scale environmental data. We evaluated the variation of species co‐occurrence patterns in climate space and estimated the influence of these patterns on the distribution of the focal species for current and projected future climates. Location Europe. Methods We used ICP Forest Level 1 data as well as climatic, topographic and edaphic variables. First, correlations between the relative abundance of European beech (Fagus sylvatica) and three major competitor species (Picea abies, Pinus sylvestris and Quercus robur) were analysed in environmental space, and then projected to geographic space. Second, a sensitivity analysis was performed using generalized additive models (GAM) to evaluate where and how much the predicted F. sylvatica distribution varied under current and future climates if potential competitor species were included or excluded. We evaluated if these areas coincide with current species co‐occurrence patterns. Results Correlation analyses supported the stress‐gradient hypothesis: towards favourable growing conditions of F. sylvatica, its abundance was strongly linked to the abundance of its competitors, while this link weakened towards unfavourable growing conditions, with stronger correlations in the south and at low elevations than in the north and at high elevations. The sensitivity analysis showed a potential spatial segregation of species with changing climate and a pronounced shift of zones where co‐occurrence patterns may play a major role. Main conclusions Our results demonstrate the importance of species co‐occurrence patterns for calibrating improved species distribution models for use in projections of climate effects. The correlation approach is able to localize European areas where inclusion of biotic predictors is effective. The climate‐induced spatial segregation of the major tree species could have ecological and economic consequences.     

Global change effects on Bromus tectorum L. (Poaceae) at its high‐elevation range margin

Global change is likely to affect invasive species distribution, especially at range margins. In the eastern Sierra Nevada, California, USA, the invasive annual grass, Bromus tectorum, is patchily distributed and its impacts have been minimal compared with other areas of the Intermountain West. We used a series of in situ field manipulations to determine how B. tectorum might respond to changing climatic conditions and increased nitrogen deposition at the high‐elevation edge of its invaded range. Over 3 years, we used snow fences to simulate changes in snowpack, irrigation to simulate increased frequency and magnitude of springtime precipitation, and added nitrogen (N) at three levels (0, 5, and 10 g m^?2) to natural patches of B. tectorum growing under the two dominant shrubs, Artemisia tridentata and Purshia tridentata, and in intershrub spaces (INTR). We found that B. tectorum seedling density in April was lower following deeper snowpack possibly due to delayed emergence, yet there was no change in spikelet production or biomass accumulation at the time of harvest. Additional spring rain events increased B. tectorum biomass and spikelet production in INTR plots only. Plants were primarily limited by water in 2009, but colimited by N and water in 2011, possibly due to differences in antecedent moisture conditions at the time of treatments. The threshold at which N had an effect varied with magnitude of water additions. Frequency of rain events was more influential than magnitude in driving B. tectorum growth and fecundity responses. Our results suggest that predicted shifts from snow to rain could facilitate expansion of B. tectorum at high elevation depending on timing of rain events and level of N deposition. We found evidence for P‐limitation at this site and an increase in P‐availability with N additions, suggesting that stoichiometric relationships may also influence B. tectorum spread.     

Silver fir stand productivity is enhanced when mixed with Norway spruce: evidence based on large‐scale inventory data and a generic modelling approach

Questions: How to evaluate the mixture effect on basal area increment in two‐species forest stands? Is a mixed Norway spruce–silver fir stand more productive than pure adjacent stands of either species? How to develop generic modelling approaches to assess mixture effects in forest stands? Location: In addition to a case study on Norway spruce–silver fir stands in French mountain forests, the generic approach used goes beyond local applications. Methods: We took advantage of National Forest Inventory data to develop a unique stand basal‐area‐increment model for pure and mixed stands of Norway spruce and silver fir that responds to ecological site conditions. The database was made up of 284 pure Norway spruce stands, 196 pure silver fir stands, and 323 mixed stands of these species. Results: Pure silver fir basal area increment is strongly influenced by spring climatic conditions, whereas pure Norway spruce is more influenced by soil conditions. The mixture of these species has a positive effect on silver fir, which decreases as the proportion of fir increases. In contrast, the mixture has no noticeable effect on Norway spruce. Conclusion: We developed a stand basal‐area‐increment model evidencing an advantage of the mixture on silver fir basal area increment, but not on Norway spruce. The mathematical formulation of the model developed is generic and can be used in all two‐species mixture situations. It also makes it possible to compare different mixture situations with each other.     

Subalpine vegetation pattern three decades after stand‐replacing fire: effects of landscape context and topography on plant community composition,tree regeneration,and diversity

Objective: Our purpose was to characterize vegetation compositional patterns, tree regeneration, and plant diversity, and their relationships to landscape context, topography, and light availability across the margins of four stand‐replacing subalpine burns. Location: Four 1977 to 1978 burns east of the Continental Divide in Colorado: the Ouzel burn, a burn near Kenosha Pass, the Badger Mountain burn, and the Maes Creek burn. Methods: Vegetation and environmental factors were sampled in 200 0.01‐ha plots on transects crossing burn edges, and stratified by elevation. We utilized dissimilarity indices, mixed‐effects models, and randomization tests to assess relationships between vegetation and environment. Results: Three decades after wildfire, plant communities exhibited pronounced compositional shifts across burn edges. Tree regeneration decreased with increasing elevation and distance into burn interiors; concomitant increases in forbs and graminoids were linked to greater light availability. Richness was roughly doubled in high‐severity burn interiors due to the persistence of a suite of native species occurring primarily in this habitat. Richness rose with distance into burns, but declined with increasing elevation. Only three of 188 plant species were non‐native; these were widespread, naturalized species that comprised <1% total cover. Conclusions: These subalpine wildfires generated considerable, persistent increases in plant species richness at local and landscape scales, and a diversity of plant communities. The findings suggest that fire suppression in such systems must lead to reduced diversity. Concerns about post‐fire invasion by exotic plants appear unwarranted in high‐elevation wilderness settings.     

Short‐ and long‐term consequences of early developmental conditions: a case study on wild and domesticated zebra finches

Divergent selection pressures among populations can result not only in significant differentiation in morphology, physiology and behaviour, but also in how these traits are related to each other, thereby driving the processes of local adaptation and speciation. In the Australian zebra finch, we investigated whether domesticated stock, bred in captivity over tens of generations, differ in their response to a life‐history manipulation, compared to birds taken directly from the wild. In a ‘common aviary’ experiment, we thereto experimentally manipulated the environmental conditions experienced by nestlings early in life by means of a brood size manipulation, and subsequently assessed its short‐ and long‐term consequences on growth, ornamentation, immune function and reproduction. As expected, we found that early environmental conditions had a marked effect on both short‐ and long‐term morphological and life‐history traits in all birds. However, although there were pronounced differences between wild and domesticated birds with respect to the absolute expression of many of these traits, which are indicative of the different selection pressures wild and domesticated birds were exposed to in the recent past, manipulated rearing conditions affected morphology and ornamentation of wild and domesticated finches in a very similar way. This suggests that despite significant differentiation between wild and domesticated birds, selection has not altered the relationships among traits. Thus, life‐history strategies and investment trade‐offs may be relatively stable and not easily altered by selection. This is a reassuring finding in the light of the widespread use of domesticated birds in studies of life‐history evolution and sexual selection, and suggests that adaptive explanations may be legitimate when referring to captive bird studies.