Pyrodiversity begets plant–pollinator community diversity

Fire has a major impact on the structure and function of many ecosystems globally. Pyrodiversity, the diversity of fires within a region (where diversity is based on fire characteristics such as extent, severity, and frequency), has been hypothesized to promote biodiversity, but changing climate and land management practices have eroded pyrodiversity. To assess whether changes in pyrodiversity will have impacts on ecological communities, we must first understand the mechanisms that might enable pyrodiversity to sustain biodiversity, and how such changes might interact with other disturbances such as drought. Focusing on plant–pollinator communities in mixed‐conifer forest with frequent fire in Yosemite National Park, California, we examine how pyrodiversity, combined with drought intensity, influences those communities. We find that pyrodiversity is positively related to the richness of the pollinators, flowering plants, and plant–pollinator interactions. On average, a 5% increase in pyrodiversity led to the gain of approximately one pollinator and one flowering plant species and nearly two interactions. We also find that a diversity of fire characteristics contributes to the spatial heterogeneity (β‐diversity) of plant and pollinator communities. Lastly, we find evidence that fire diversity buffers pollinator communities against the effects of drought‐induced floral resource scarcity. Fire diversity is thus important for the maintenance of flowering plant and pollinator diversity and predicted shifts in fire regimes to include less pyrodiversity compounded with increasing drought occurrence will negatively influence the richness of these communities in this and other forested ecosystems. In addition, lower heterogeneity of fire severity may act to reduce spatial turnover of plant–pollinator communities. The heterogeneity of community composition is a primary determinant of the total species diversity present in a landscape, and thus, lower pyrodiversity may negatively affect the richness of plant–pollinator communities across large spatial scales.     

Landscape‐scale effects of fire on bird assemblages: does pyrodiversity beget biodiversity?

Aim A common strategy for conserving biodiversity in fire‐prone environments is to maintain a diversity of post‐fire age classes at the landscape scale, under the assumption that ‘pyrodiversity begets biodiversity’. Another strategy is to maintain extensive areas of a particular seral state regarded as vital for the persistence of threatened species, under the assumption that this will also cater for the habitat needs of other species. We investigated the likely effects of these strategies on bird assemblages in tree mallee vegetation, characterized by multi‐stemmed Eucalyptus species, where both strategies are currently employed. Location The semi‐arid Murray Mallee region of south‐eastern Australia. Methods We systematically surveyed birds in 26 landscapes (each 4‐km diameter), selected to represent gradients in the diversity of fire age classes and the proportion of older vegetation (> 35 years since fire). Additional variables were measured to represent underlying vegetation‐ or fire‐mediated properties of the landscape, as well as its biogeographic context. We used an information‐theoretic approach to investigate the relationships between these predictor variables and the species richness of birds (total species, threatened species and rare species). Results Species richness of birds was not strongly associated with fire‐mediated heterogeneity. Species richness was associated with increasing amounts of older vegetation in landscapes, but not with the proportion of recently burned vegetation in landscapes. Main conclusions The preference of many mallee birds for older vegetation highlights the risk of a blanket application of the ‘pyrodiversity begets biodiversity’ paradigm. If application of this paradigm involved converting large areas from long unburned to recently burned vegetation to increase fire‐mediated heterogeneity in tree mallee landscapes, our findings suggest that this could threaten birds. This research highlights the value of adopting a landscape‐scale perspective when evaluating the utility of fire‐management strategies intended to benefit biodiversity.     

Pyrodiversity interacts with rainfall to increase bird and mammal richness in African savannas

Fire is a fundamental process in savannas and is widely used for management. Pyrodiversity, variation in local fire characteristics, has been proposed as a driver of biodiversity although empirical evidence is equivocal. Using a new measure of pyrodiversity (Hempson et al.), we undertook the first continent‐wide assessment of how pyrodiversity affects biodiversity in protected areas across African savannas. The influence of pyrodiversity on bird and mammal species richness varied with rainfall: strongest support for a positive effect occurred in wet savannas (> 650 mm/year), where species richness increased by 27% for mammals and 40% for birds in the most pyrodiverse regions. Range‐restricted birds were most increased by pyrodiversity, suggesting the diversity of fire regimes increases the availability of rare niches. Our findings are significant because they explain the conflicting results found in previous studies of savannas. We argue that managing savanna landscapes to increase pyrodiversity is especially important in wet savannas.     

Conservation value of low fire frequency in tropical savannas: Ants in monsoonal northern Australia

The conservation values of ‘old‐growth’ forests in landscapes subject to repeated disturbance by fire or logging have received considerable conservation attention. However, little is known of the conservation values of old‐growth sites in ecosystems with an evolutionary history of highly frequent disturbance. Here we address the value of low fire frequency (<1 fire/10 years) in tropical savannas, the world's most fire‐prone biome, in terms of ant biodiversity. We do this by comparing savanna ant communities within the Territory Wildlife Park (TWP) near Darwin in the Australian monsoonal tropics, which has experienced a low incidence of fire over 25 years due to active fire exclusion, with those of adjacent (outside) sites experiencing the ambient fire regime of burning every 2–5 years. Ants were sampled using terrestrial and arboreal pitfall traps at 16 sites, eight each inside and outside TWP. More than 16 000 ants were recorded during the study, representing a total of 98 ant species from 30 genera. More species in total were recorded outside (90) than inside (74) TWP, but there was no difference in mean site species richness or abundance, and overall species composition was similar. All species recorded inside TWP are common and widespread throughout the savanna landscapes of the broader region, in the absence of active fire exclusion. Low fire frequency at the Territory Wildlife Park therefore does not appear to have enhanced regional ant conservation values. Our findings reinforce the importance of targeting fire regimes that are clearly linked to positive conservation outcomes, rather than assuming a need for maximum ‘pyrodiversity’.     

Continent‐level drivers of African pyrodiversity

Pyrodiversity, which describes fire variability over space and time, is believed to increase habitat heterogeneity and thereby promote biodiversity. However, to date there is no standardised metric for quantifying pyrodiversity, and so broad geographic patterns and drivers of pyrodiversity remain unexplored. We present the first generalizable method to quantify pyrodiversity, and use it to address the fundamental questions of what drives pyrodiversity, which fire attributes constrain pyrodiversity under different conditions, and whether pyrodiversity is spatial grain‐dependent. We linked the MODIS burned area and active fire products to measure fire size, seasonal timing, return interval, and intensity for 2.2 million individual fires in sub‐Saharan Africa from 2000–2015. We then quantified pyrodiversity as a four‐dimensional hypervolume described by fire attributes within a grid cell, for any spatial grain of analysis. Environmental (rainfall, vegetation, soils, and topography) and human‐associated (cattle biomass, cropland area, and human population density) variables were assessed as potential drivers of pyrodiversity. Rainfall was the main environmental driver of pyrodiversity, with higher pyrodiversity in drier regions (< 650 mm yr^–1). Pyrodiversity was not strongly associated with human‐associated variables across Africa. Rainfall and a human influence index had clear but contrasting effects on the variability of fire size, seasonal timing, return interval, and intensity. Our analyses show that fire size and seasonal timing constrain pyrodiversity in wetter regions, whereas none of the fire attributes pose a strong constraint in drier regions. We found no evidence that pyrodiversity was spatial grain‐dependent when recalculated at 5‐minute grain increments from 15 to 120 minutes. We hypothesise that the strongest positive effect of pyrodiversity on biodiversity in all its forms will occur at intermediate precipitation (650–1300 mm yr^–1), where fire plays an important role in shaping vegetation structure and where pyrodiversity is still quite high.     

Atlantic floodplain meadows: influence of hydrological gradients and management on sciomyzid (Diptera) assemblages

Maintaining biodiversity is central to maintaining ecosystem functionality of wetlands. Hydrology has the strongest influence on wetland biodiversity, second to which agriculture is the most influential factor. This study investigates the influence of hydrology and farming practices on the abundance, species richness and composition of dipteran communities on temperate Atlantic floodplain hay meadows. Insects were sampled by sweep-net across twenty-four vegetation zones for which hydrological variables were calculated by combining river level data with fine-scale topographical data. Plant communities were surveyed using relevés and land owners were interviewed to gather data on current and past management regimes. A total of twenty-two sciomyzid species were recorded; over one-third of the Irish fauna. Flood depth and duration were found to have the strongest influence on sciomyzids, syrphids and plants. Sciomyzid species richness and total abundance were both positively correlated with hydroperiod and flood depth while both plants and syrphids responded negatively to increases. The difference in response highlights the need to assess more than one taxonomic group, when assessing the impact of changing environmental variables on biodiversity. Whereas vegetation structure drives changes in sciomyzid indicator species, plant species richness and composition, past management regimes and current nutrient inputs do not appear to influence these species. Thus, while the maintenance of the hydrological heterogeneity and the diversity of mowing regimes is important in maintaining biodiversity, variation in nutrient inputs and previous management (at least within the range here investigated) is likely to be of lesser importance for Syrphidae and Sciomyzidae.     

Stochastic losses of fire‐dependent endemic herbs revealed by a 65‐year chronosequence of dispersal‐limited woody plant encroachment

The factors responsible for maintaining diverse groundcover plant communities of high conservation value in frequently burned wet pine savannas are poorly understood. While most management involves manipulating extrinsic factors important in maintaining species diversity (e.g., fire regimes), most ecological theory (e.g., niche theory and neutral theory) examines how traits exhibited by the species promote species coexistence. Furthermore, although many ecologists focus on processes that maintain local species diversity, conservation biologists have argued that other indices (e.g., phylogenetic diversity) are better for evaluating assemblages in terms of their conservation value. I used a null model that employed beta‐diversity calculations based on Raup–Crick distances to test for deterministic herbaceous species losses associated with a 65‐year chronosequence of woody species encroachment within each of three localities. I quantified conservation value of assemblages by measuring taxonomic distinctness, endemism, and floristic quality of plots with and without woody encroachment. Reductions in herb species richness per plot attributable to woody encroachment were largely stochastic, as indicated by a lack of change in the mean or variance in beta‐diversity caused by woody encroachment in the savannas studied here. Taxonomic distinctness, endemism, and floristic quality (when summed across all species) were all greater in areas that had not experienced woody encroachment. However, when corrected for local species richness, only average endemism and floristic quality of assemblages inclusive of herbs and woody plants were greater in areas that had not experienced woody encroachment, due to the more restricted ranges and habitat requirements of herbs. Results suggest that frequent fires maintain diverse assemblages of fire‐dependent herb species endemic to the region. The stochastic loss of plant species, irrespective of their taxonomic distinctness, to woody encroachment suggests that the relevance of niche partitioning or phylogenetic diversity to the management of biodiversity in wet pine savannas is minimal.     

Relative roles of ecological and energetic constraints,diversification rates and region history on global species richness gradients

Regions worldwide differ markedly in species richness. Here, for birds and mammals worldwide, we directly compare four sets of hypotheses regarding geographical richness gradients: (1) evolutionary, emphasising heterogeneity in diversification rates, (2) historical, related to differences in region ages and sizes, (3) energetic, associated with variation in productive or ambient energy and (4) ecological, reflecting differences in ecological niche diversity. Among highly independent regions, or ‘evolutionary arenas’, we find that richness is weakly influenced by richness‐standardised ecological niche diversity, questioning the significance of ecological constraints for producing large‐scale diversity gradients. In contrast, we find strong evidence for the importance of region area and its changes over time, together with a role for temperature. These predictors affect richness predominately directly without concomitant positive effects on diversification rates. This suggests that regional richness is governed by historical and evolutionary processes, which promote region‐specific accumulation of diversity through time or following asymmetrical dispersal.     

Into the Andes: multiple independent colonizations drive montane diversity in the Neotropical clearwing butterflies Godyridina

Understanding why species richness peaks along the Andes is a fundamental question in the study of Neotropical biodiversity. Several biogeographic and diversification scenarios have been proposed in the literature, but there is confusion about the processes underlying each scenario, and assessing their relative contribution is not straightforward. Here, we propose to refine these scenarios into a framework which evaluates four evolutionary mechanisms: higher speciation rate in the Andes, lower extinction rates in the Andes, older colonization times and higher colonization rates of the Andes from adjacent areas. We apply this framework to a species‐rich subtribe of Neotropical butterflies whose diversity peaks in the Andes, the Godyridina (Nymphalidae: Ithomiini). We generated a time‐calibrated phylogeny of the Godyridina and fitted time‐dependent diversification models. Using trait‐dependent diversification models and ancestral state reconstruction methods we then compared different biogeographic scenarios. We found strong evidence that the rates of colonization into the Andes were higher than the other way round. Those colonizations and the subsequent local diversification at equal rates in the Andes and in non‐Andean regions mechanically increased the species richness of Andean regions compared to that of non‐Andean regions (‘species‐attractor’ hypothesis). We also found support for increasing speciation rates associated with Andean lineages. Our work highlights the importance of the Andean slopes in repeatedly attracting non‐Andean lineages, most likely as a result of the diversity of habitats and/or host plants. Applying this analytical framework to other clades will bring important insights into the evolutionary mechanisms underlying the most species‐rich biodiversity hotspot on the planet.     

Pyrodiversity—why managing fire in food webs is relevant to restoration ecology

The manipulation of landscape fire to maintain biodiverse, self‐sustaining ecosystems in flammable landscapes is rarely considered by restoration ecologists. Fire regimes can interact with ecological processes, food webs, and biodiversity in complex ways (here called pyrodiversity) and understanding these complexities could be used to promote restoration and resilience. We illustrate this using an example from northern Australia. Understanding and using pyrodiversity in ecological restoration programs will be intellectually and financially challenging. In Australia, the considerable technical and financial resources of the mining industry could support such restoration programs, yet redirecting these resources from the current narrow focus on restoring native vegetation cover at the mine‐affected site requires overcoming entrenched attitudes among policymakers and restoration ecologists.     

Plant diversity,biogeography and environment in Iberia: Patterns and possible causal factors

Abstract. We associated patterns of plant diversity with possible causal factors by considering 93 local regions in the Iberian Peninsula and Balearic Islands with respect to biogeography, environmental favourability, and environmental heterogeneity, and their relationship with measured species diversity at four different scales: mean local species richness standardized at a grain of 100 m², total species richness in a community type within a region (regional community richness), mean compositional similarity, and mosaic diversity. Local regions in biogeographic transition zones to the North African and Atlantic floras had higher regional community richness and greater mosaic diversity than did non‐transitional regions, whereas no differences existed in mean local species richness or mean compositional similarity. Mean local species richness was positively related to environmental favourability as measured by actual evapotranspiration, but negatively related to total precipitation and temporal heterogeneity in precipitation. Mean local species richness was greatest in annual grassland and dwarf shrubland communities, and on calcareous bedrock types. Regional community richness was similarly related to actual evapotranspiration and total precipitation, but in addition was positively related to spatial heterogeneity in topography and soil water holding capacity. Mean compositional similarity decreased with increasing spatial heterogeneity and temperature seasonality. Mosaic diversity, a measure of complexity, increased with increasing local and regional richness. We hypothesize that these relationships can be explained by four ecological and evolutionary classes of causal factors: numbers of individuals, intermediate environments, limits to adaptation, and niche variation. These factors operate at various scales and manifest themselves in various ways. For example, at the site level, apparently processes that increase the number of individuals increase mean local species richness, but at the level of the entire region no such effects were found.     

Diversity of plant evolutionary lineages promotes arthropod diversity

Large‐scale habitat destruction and climate change result in the non‐random loss of evolutionary lineages, reducing the amount of evolutionary history represented in ecological communities. Yet, we have limited understanding of the consequences of evolutionary history on the structure of food webs and the services provided by biological communities. Drawing on 11 years of data from a long‐term plant diversity experiment, we show that evolutionary history of plant communities – measured as phylogenetic diversity – strongly predicts diversity and abundance of herbivorous and predatory arthropods. Effects of plant species richness on arthropods become stronger when phylogenetic diversity is high. Plant phylogenetic diversity explains predator and parasitoid richness as strongly as it does herbivore richness. Our findings indicate that accounting for evolutionary relationships is critical to understanding the severity of species loss for food webs and ecosystems, and for developing conservation and restoration policies.     

The impact of shifts in marine biodiversity hotspots on patterns of range evolution: Evidence from the Holocentridae (squirrelfishes and soldierfishes)

One of the most striking biodiversity patterns is the uneven distribution of marine species richness, with species diversity in the Indo-Australian Archipelago (IAA) exceeding all other areas. However, the IAA formed fairly recently, and marine biodiversity hotspots have shifted across nearly half the globe since the Paleogene. Understanding how lineages have responded to shifting biodiversity hotspots represents a necessary historic perspective on the formation and maintenance of global marine biodiversity. Such evolutionary inferences are often challenged by a lack of fossil evidence that provide insights into historic patterns of abundance and diversity. The greatest diversity of squirrelfishes and soldierfishes (Holocentridae) is in the IAA, yet these fishes also represent some of the most numerous fossil taxa in deposits of the former West Tethyan biodiversity hotspot. We reconstruct the pattern of holocentrid range evolution using time-calibrated phylogenies that include most living species and several fossil lineages, demonstrating the importance of including fossil species as terminal taxa in ancestral area reconstructions. Holocentrids exhibit increased range fragmentation following the West Tethyan hotspot collapse. However, rather than originating within the emerging IAA hotspot, the IAA has acted as a reservoir for holocentrid diversity that originated in adjacent regions over deep evolutionary time scales.     

Environmental causes for plant biodiversity gradients

One of the most pervasive patterns observed in biodiversity studies is the tendency for species richness to decline towards the poles. One possible explanation is that high levels of environmental energy promote higher species richness nearer the equator. Energy input may set a limit to the number of species that can coexist in an area or alternatively may influence evolutionary rates. Within flowering plants (angiosperms), families exposed to a high energy load tend to be both more species rich and possess faster evolutionary rates, although there is no evidence that one drives the other. Specific environmental effects are likely to vary among lineages, reflecting the interaction between biological traits and environmental conditions in which they are found. One example of this is demonstrated by the high species richness of the iris family (Iridaceae) in the Cape of South Africa, a likely product of biological traits associated with reproductive isolation and the steep ecological and climatic gradients of the region. Within any set of conditions some lineages will tend to be favoured over others; however, the identity of these lineages will fluctuate with a changing environment, explaining the highly labile nature of diversification rates observed among major lineages of flowering plants.     

Plant species diversity and grazing in the Scandinavian mountains - patterns and processes at different spatial scales

There is a long tradition of grazing by semi‐domestic reindeer and sheep in alpine and sub‐alpine Scandinavian habitats, but present management regimes are questioned from a conservation point of view. In this review we discuss plant diversity patterns in the Scandinavian mountains in a global, regional and local perspective. The main objective was to identify processes that influence diversity at different spatial scales with a particular focus on grazing. In a global perspective the species pool of the Scandinavian mountains is limited. partly reflecting the general latitudinal decline of species but also historical and ecological factors operating after the latest glaciation. At the local scale, both productivity and disturbance are primary factors structuring diversity, but abiotic factors such as soil pH, snow distribution and temperature are also important. Although evidence is scarce, grazing favours local species richness in productive habitats, whereas species richness decreases with grazing when productivity is low. Regional patterns of plant diversity is set by, 1) the species pool. 2) the heterogeneity and fragmentation of communities, and 3) local diversity of each plant community. We suggest that local shifts in community composition depend both on the local grazing frequency and the return‐time of the plant community after a grazing session. In addition, an increasing number of grazing‐modified local patches homogenises the vegetation and is likely to reduce the regional plant diversity. The time scale of local shifts in community composition depends on plant colonisation and persistence, From a mechanistic point of view, diversity patterns at a regional scale also depend on the regional dynamics of single species. Colonisation is usually a slow and irregular process in alpine environments, whereas the capacity for extended local persistence is generally high. Although the poor knowledge of plant regional dynamics restricts our understanding of how grazing influences plant diversity, we conclude that grazing is a key process for maintaining biodiversity in the Scandinavian mountains.     

Ecological processes: A key element in strategies for nature conservation

Summary A common approach to nature conservation is to identify and protect natural ‘assets’ such as ecosystems and threatened species. While such actions are essential, protection of assets will not be effective unless the ecological processes that sustain them are maintained. Here, we consider the role of ecological processes and the complementary perspective for conservation arising from an emphasis on process. Many kinds of ecological processes sustain biodiversity: including climatic processes, primary productivity, hydrological processes, formation of biophysical habitats, interactions between species, movements of organisms and natural disturbance regimes. Anthropogenic threats to conservation exert their influence by modifying or disrupting these processes. Such threats extend across tenures, they frequently occur offsite, they commonly induce non‐linear responses, changes may be irreversible and the full consequences may not be experienced for lengthy periods. While many managers acknowledge these considerations in principle, there is much scope for greater recognition of ecological processes in nature conservation and greater emphasis on long time‐frames and large spatial scales in conservation planning. Practical measures that promote ecological processes include: monitoring to determine the trajectory and rate of processes; incorporating surrogates for processes in conservation and restoration projects; specific interventions to manipulate and restore processes; and planning for the ecological future before options are foreclosed. The long‐term conservation of biodiversity and the well‐being of human society depend upon both the protection of natural assets and maintaining the integrity of the ecological processes that sustain them.     

When can refuges mediate the genetic effects of fire regimes? A simulation study of the effects of topography and weather on neutral and adaptive genetic diversity in fire‐prone landscapes

Understanding how landscape heterogeneity mediates the effects of fire on biodiversity is increasingly important under global changes in fire regimes. We used a simulation experiment to investigate how fire regimes interact with topography and weather to shape neutral and selection‐driven genetic diversity under alternative dispersal scenarios, and to explore the conditions under which microrefuges can maintain genetic diversity of populations exposed to recurrent fire. Spatial heterogeneity in simulated fire frequency occurred in topographically complex landscapes, with fire refuges and fire‐prone “hotspots” apparent. Interannual weather variability reduced the effect of topography on fire patterns, with refuges less apparent under high weather variability. Neutral genetic diversity was correlated with long‐term fire frequency under spatially heterogeneous fire regimes, being higher in fire refuges than fire‐prone areas, except under high dispersal or low fire severity (low mortality). This generated different spatial genetic structures in fire‐prone and fire‐refuge components of the landscape, despite similar dispersal. In contrast, genetic diversity was only associated with time since the most recent fire in flat landscapes without predictable refuges and hotspots. Genetic effects of selection driven by fire‐related conditions depended on selection pressure, migration distance and spatial heterogeneity in fire regimes. Allele frequencies at a locus conferring higher fitness under successional environmental conditions followed a pattern of “temporal adaptation” to contemporary conditions under strong selection pressure and high migration. However, selected allele frequencies were correlated with spatial variation in long‐term mean fire frequency (relating to environmental predictability) under weak dispersal, low selection pressure and strong spatial heterogeneity in fire regimes.     

Trait diversity on the phylogeny of cerrado woody species

During the Neogene, the Brazilian cerrado became established as a large‐scale vegetation type. Cerrado lineages started to diversify less than 10 million years ago, coinciding with the rise to dominance of flammable C₄ grasses and the expansion of the savanna biome. Cerrado lineages are strongly associated with adaptations to fire and have sister groups in fire‐free nearby forests, implying that the cerrado formed in situ via adaptive shifts to resist fire. By including phylogeny into the analysis of biological traits, we investigated trait diversity of cerrado woody species in a phylogenetic context, sampling a cerrado site in central Brazil. Decomposing trait diversity along the nodes of a phylogenetic tree of cerrado woody species, we found that the rate of trait diversification was higher in the past, coinciding with the major species diversification of angiosperms in mid‐Cretaceous, long before the cerrado originated. Some more recent adaptive shifts to resist fire, however, must have occurred during the origin and expansion of the cerrado woody flora. Analysing values of each trait separately at the tips of the phylogenetic tree, we found that most trait values were randomly distributed, probably because we analysed only species that had already been filtered by drought, fire, and soil. Analysing values of all traits simultaneously at the tips, we found close to root events and broad, macro‐evolutionary patterns, called ‘global structures’, opposing some lineages, especially Fabaceae and Myrtaceae, with different ecological strategies. Fabaceae presented compound, large, tender leaves, with high nitrogen content due to symbiosis with nitrogen‐fixing bacteria, and Myrtaceae presented simple, small, tough leaves, with low nitrogen and high potassium content. We also found relatively recent events that induced divergence of the evolutionary strategies close to the tips, called ‘local structures’, involving more recent changes in most lineages.     

Advancing the metabolic theory of biodiversity

A component of metabolic scaling theory has worked towards understanding the influence of metabolism over the generation and maintenance of biodiversity. Specific models within this ‘metabolic theory of biodiversity’ (MTB) have addressed temperature gradients in speciation rate and species richness, but the scope of MTB has been questioned because of empirical departures from model predictions. In this study, we first show that a generalized MTB is not inconsistent with empirical patterns and subsequently implement an eco‐evolutionary MTB which has thus far only been discussed qualitatively. More specifically, we combine a functional trait (body mass) approach and an environmental gradient (temperature) with a dynamic eco‐evolutionary model that builds on the current MTB. Our approach uniquely accounts for feedbacks between ecological interactions (size‐dependent competition and predation) and evolutionary rates (speciation and extinction). We investigate a simple example in which temperature influences mutation rate, and show that this single effect leads to dynamic temperature gradients in macroevolutionary rates and community structure. Early in community evolution, temperature strongly influences speciation and both speciation and extinction strongly influence species richness. Through time, niche structure evolves, speciation and extinction rates fall, and species richness becomes increasingly independent of temperature. However, significant temperature‐richness gradients may persist within emergent functional (trophic) groups, especially when niche breadths are wide. Thus, there is a strong signal of both history and ecological interactions on patterns of species richness across temperature gradients. More generally, the successful implementation of an eco‐evolutionary MTB opens the perspective that a process‐based MTB can continue to emerge through further development of metabolic models that are explicit in terms of functional traits and environmental gradients.     

Pyrodiversity promotes interaction complementarity and population resistance

Theory predicts that network characteristics may help anticipate how populations and communities respond to extreme climatic events, but local environmental context may also influence responses to extreme events. For example, altered fire regimes in many ecosystems may significantly affect the context for how species and communities respond to changing climate. In this study, I tested whether the responses of a pollinator community to extreme drought were influenced by the surrounding diversity of fire histories (pyrodiversity) which can influence their interaction networks via changing partner availability. I found that at the community level, pyrodiverse landscapes promote functional complementarity and generalization, but did not consistently enhance functional redundancy or resistance to simulated co‐extinction cascades. Pyrodiversity instead supported flexible behaviors that enable populations to resist perturbations. Specifically, pollinators that can shift partners and network niches are better able to take advantage of the heterogeneity generated by pyrodiversity, thereby buffering pollinator populations against changes in plant abundances. These findings suggest that pyrodiversity is unlikely to improve community‐level resistance to droughts, but instead promotes population resistance and community functionality. This study provides unique evidence that resistance to extreme climatic events depends on both network properties and historical environmental context.