Lower thermal tolerance in nocturnal than in diurnal ants: a challenge for nocturnal ectotherms facing global warming

1. The thermal adaptation hypothesis proposes that because thermoregulation involves a high metabolic cost, thermal limits of organisms must be locally adapted to temperatures experienced in their environments. There is evidence that tolerance to high temperatures decreases in insects inhabiting colder habitats and microclimates. However, it is not clear if thermal limits of ectotherms with contrasting temporal regimes, such as diurnal and nocturnal insects, are also adapted to temperatures associated with their circadian activities. 2. This study explores differences in heat tolerance among diurnal and nocturnal ant species in four ecosystems in Mexico: tropical montane, tropical rainforest, subtropical dry forests, and high‐elevation semi‐desert. 3. The critical thermal maximum (CT_max), i.e. the temperature at which ants lost motor control, was estimated for diurnal and nocturnal species. CT_max for 19 diurnal and 12 nocturnal ant species distributed among 45 populations was also estimated. 4. Semi‐desert and subtropical dry forest ants displayed higher tolerances to high temperatures than did ants in tropical rainforest. The lowest tolerance to high temperatures was recorded in tropical montane forest ants. In general, among all habitats, the CT_max of nocturnal ants was lower than that of diurnal ants. 5. An increase in nocturnal temperatures, combined with lower tolerance to high temperatures, may represent a substantial challenge for nocturnal ectotherms in a warming world.     

Thermal adaptation generates a diversity of thermal limits in a rainforest ant community

The Thermal Adaptation Hypothesis posits that the warmer, aseasonal tropics generates populations with higher and narrower thermal limits. It has largely been tested among populations across latitudes. However, considerable thermal heterogeneity exists within ecosystems: across 31 trees in a Panama rainforest, surfaces exposed to sun were 8 °C warmer and varied more in temperature than surfaces in the litter below. Tiny ectotherms are confined to surfaces and are variously submerged in these superheated boundary layer environments. We quantified the surface CT_min and CT_maxs (surface temperatures at which individuals grew torpid and lost motor control, respectively) of 88 ant species from this forest; they ranged in average mass from 0.01 to 57 mg. Larger ants had broader thermal tolerances. Then, for 26 of these species we again tested body CT_maxs using a thermal dry bath to eliminate boundary layer effects: body size correlations observed previously disappeared. In both experiments, consistent with Thermal Adaptation, CT_maxs of canopy ants averaged 3.5–5 °C higher than populations that nested in the shade of the understory. We impaled thermocouples in taxidermy mounts to further quantify the factors shaping operative temperatures for four ant species representing the top third (1–30 mg) of the size distribution. Extrapolations suggest the smallest 2/3rds of species reach thermal equilibrium in <10s. Moreover, the large ants that walk above the convective superheated surface air also showed more net heating by solar radiation, with operative temperatures up to 4 °C higher than surrounding air. The thermal environments of this Panama rainforest generate a range of CT_max subsuming 74% of those previously recorded for ant populations worldwide. The Thermal Adaptation Hypothesis can be a powerful tool in predicting diversity of thermal limits within communities. Boundary layer temperatures are likely key to predicting the future of Earth's tiny terrestrial ectotherm populations.     

Species interactions and thermal constraints on ant community structure

Patterns of species occurrence and abundance are influenced by abiotic factors and biotic interactions, but these factors are difficult to disentangle without experimental manipulations. In this study, we used observational and experimental approaches to investigate the role of temperature and interspecific competition in controlling the structure of ground‐foraging ant communities in forests of the Siskiyou Mountains of southwestern Oregon. To assess the potential role of competition, we first used null model analyses to ask whether species partition temporal and/or spatial environments. To understand how thermal tolerances influence the structure of communities, we conducted a laboratory experiment to estimate the maximum thermal tolerance of workers and a field experiment in which we added shaded microhabitats and monitored the response of foragers. Finally, to evaluate the roles of temperature and interspecific competition in the field, we simultaneously manipulated shading and the presence of a dominant competitor (Formica moki). The foraging activity of species broadly overlapped during the diurnal range of temperatures. Species co‐occurrence patterns varied across the diurnal temperature range: species were spatially segregated at bait stations at low temperatures, but co‐occurred randomly at high temperatures. The decreased abundance of the co‐occurring thermophilic Temnothorax nevadensis in shaded plots was a direct effect of shading and not an indirect effect of competitive interactions. Thermal tolerance predicted the response of ant species to the shading experiment: species with the lowest tolerances to high temperatures showed the greatest increase in abundance in the shaded plots. Moreover, species with more similar thermal tolerance values segregated more frequently on baits than did species that differed in their thermal tolerances. Collectively, our results suggest that thermal tolerances of ants may mediate competitive effects in habitats that experience strong diurnal temperature fluctuations.     

Alternative adaptations, sympatric speciation and the evolution of parasitic, inquiline ants

Inquiline ant species are workerless social parasites whose queens reproduce in colonies of other species alongside the host queens. Inquilines arise either when one non-parasitic species evolves into an inquiline parasite of another non-parasitic species (the interspecific hypothesis), or by the speciation of intraspecific inquilines from their host stock (the intraspecific hypothesis): it is unlikely that inquilines evolve from other forms of social parasite. This paper reviews the evidence for and against the inter-and intraspecific hypotheses. All inquilines are close phylogenetic relatives of their host species (loose ‘Emery's rule’), and some are their host's closest relative (strict ‘Emery's rule’). A problem for the interspecific hypothesis is how to explain the strict Emery's rule, because phylogenetic constraints on host choice are probably quite weak. By contrast, the intraspecific hypothesis has difficulty accounting for the parasites' sympatric reproductive isolation. Facultative polygyny, in which queens may found colonies alone or by adoption into an existing multi-queen colony, should promote the evolution of small intraspecific inquilines. This is because small colony-founding queens should preferentially seek adoption, which provides the opportunity to produce a sexual-only brood. We suggest that microgynes, i.e. miniature queens found in some polygynous ants, represent such parasites. We review the evidence that inquiline species have evolved intraspecifically from microgynes in Myrmica ants. The coexistence within a species of a monogynous (singly-queened) and a polygynous form is probably a phenomenon usually unconnected with inquiline evolution. The reproductive isolation of intraspecific inquilines plausibly arises from divergent breeding behaviour associated with the parasites' small size. Such divergence could involve either a temporal separation in mating episodes, with small parasites maturing early, or a spatial separation, with small males being sexually-selected to mate near the nest with small queens seeking adoption, instead of in mating aggregations. We conclude that inquiline species strictly following Emery's rule could have evolved by the intraspecific route. If so, such species provide evidence for West-Eberhard's “alternative adaptation” hypothesis that between-species diversity frequently stems from diversity within species. They also represent likely cases of sympatric speciation. We suggest work on the parasites' phytogeny, genetics, behaviour and mating biology to test these conclusions further.     

Nocturnal resource defence in aphid‐tending ants of northern Patagonia

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Pattern of ant diversity in Korea: An empirical test of Rapoport's altitudinal rule

Two diversity patterns (hump-shaped and monotonic decrease) frequently occur along altitude or latitude gradients. We examined whether patterns of ant species richness along altitudes in South Korea can be described by these patterns and whether ranges of ant species follow Rapoport's altitudinal rule. Ants on 12 high mountains (> 1100 m) throughout South Korea (from 33° N to 38° N) were surveyed using pitfall traps at intervals of 200–300 m altitude. The temperatures at the sampling sites were determined from digital climate maps. Ant species richness decreased monotonically along the altitudinal gradient and increased along the temperature gradient. However, species richness of cold-adapted species (highland species) showed a hump-shaped pattern along altitude and temperature gradients. The altitude and temperature ranges of ant species followed Rapoport's rule. Sampling site temperature ranges were significantly correlated with coldness. Therefore, Rapoport's rule can be explained by high cold-tolerance of species inhabiting high altitudes or latitudes.     

Myrmica rubra microhabitat selection and putative ecological impact

1. Myrmica rubra (European fire ant) has invaded northern latitude coastal areas in North America. This macroscale distribution suggests that M. rubra is moisture‐ and temperature‐limited, but the distribution of the invaded range may reflect the legacy of original introduction locations preserved by limited dispersal. 2. This study examined a two‐decade population change in M. rubra (1994–2015) and the microscale abiotic (moisture and temperature), biotic (plants), anthropogenic (pesticide) and physiological (thermal tolerance) limits on the invasion at the Tifft Nature Preserve in Buffalo, NY (U.S.A.). Changes in the abundance of native ants and other invertebrates were also examined. 3. Despite localised declines with pesticide treatments, overall M. rubra forager abundance increased 27% between 1994 and 2015. Abundance increased the most in the warmest areas (native ants were unaffected by temperature), but M. rubra colony locations were strongly linked to higher soil moisture and lower soil temperature. Myrmica rubra ants also exhibited relatively low thermal tolerances consistent with high‐latitude and high‐elevation ants. 4. Where local M. rubra populations increased the most, native ant species decreased, and where local M. rubra populations declined, native ant species increased. Some arthropod species had lower abundance with M. rubra presence, but the impacts were less striking. 5. Myrmica rubra population growth was promoted at the microhabitat scale where relatively higher temperatures prompted foraging, and relatively lower temperatures and high moisture supported nesting. These results suggest that macroscale M. rubra invaded‐range distributions in northern climates near coastal areas are replicated at the microscale where the ant prefers cooler, moister microsites.     

Upward ant distribution shift corresponds with minimum,not maximum,temperature tolerance

Rapid climate change may prompt species distribution shifts upward and poleward, but species movement in itself is not sufficient to establish climate causation. Other dynamics, such as disturbance history, may prompt species distribution shifts resembling those expected from rapid climate change. Links between species distributions, regional climate trends and physiological mechanism are needed to convincingly establish climate‐induced species shifts. We examine a 38‐year shift (1974–2012) in an elevation ecotone between two closely related ant species, Aphaenogaster picea and A. rudis. Even though A. picea and A. rudis are closely related with North American distributions that sometimes overlap, they also exhibit local‐ and regional‐scale differences in temperature requirements so that A. rudis is more southerly and inhabits lower elevations whereas A. picea is more northerly and inhabits high elevations. We find considerable movement by the warm‐habitat species upward in elevation between 1974 and 2012 with A. rudis, replacing the cold‐habitat species, A. picea, along the southern edge of the Appalachian Mountain chain in north Georgia, USA. Concomitant with the distribution shifts, regional mean and maximum temperatures remain steady (1974–2012), but minimum temperatures increase. We collect individuals from the study sites and subject them to thermal tolerance testing in a controlled setting and find that maximum and minimum temperature acclimatization occurs along the elevation gradient in both species, but A. rudis consistently becomes physiologically incapacitated at minimum and maximum temperatures 2 °C higher than A. picea. These results indicate that rising minimum temperatures allow A. rudis to move upward in elevation and displace A. picea. Given that Aphaenogaster ants are the dominant woodland seed dispersers in eastern deciduous forests, and that their thermal tolerances drive distinct differences in temperature‐cued synchrony with early blooming plants, these climate responses not only impact ant‐ant interactions, but might have wide implications for ant‐plant interactions.     

Who likes it hot? A global analysis of the climatic,ecological, and evolutionary determinants of warming tolerance in ants

Effects of climate warming on wild populations of organisms are expected to be greatest at higher latitudes, paralleling greater anticipated increases in temperature in these regions. Yet, these expectations assume that populations in different regions are equally susceptible to the effects of warming. This is unlikely to be the case. Here, we develop a series of predictive models for physiological thermal tolerances in ants based on current and future climates. We found that tropical ants have lower warming tolerances, a metric of susceptibility to climate warming, than temperate ants despite greater increases in temperature at higher latitudes. Using climatic, ecological and phylogenetic data, we refine our predictions of which ants (across all regions) were most susceptible to climate warming. We found that ants occupying warmer and more mesic forested habitats at lower elevations are the most physiologically susceptible to deleterious effects of climate warming. Phylogenetic history was also a strong indicator of physiological susceptibility. In short, we find that ants that live in the canopies of hot, tropical forest are the most at risk, globally, from climate warming. Unfortunately this is where many, perhaps most, ant and other species on Earth live.     

Mountain passes are higher not only in the tropics

A recent article in Ecography by Zuloaga and Kerr (2016) addressed a prediction of Janzen's (1967) classic contention that mountain passes are higher in the tropics: species assemblages separated by steep thermal gradients are less similar than assemblages separated by small thermal gradients. Their results have some surprising and important additional implications. In the New World, mountain passes are in fact higher in both the tropics and near 55°N. This fact allows a strong test of Simpson's (1964) hypothesis that thermal barriers promote allopatric speciation, which leads to higher species richness. Combined with the results of Zuloaga and Kerr, the Simpson/Janzen hypothesis predicts a peak of richness in southern Alaskan and northern British Columbia. Published data are clearly inconsistent with this prediction. High thermal barriers to dispersal do not necessarily lead to greater species richness.     

Can temperate insects take the heat? A case study of the physiological and behavioural responses in a common ant, Iridomyrmex purpureus (Formicidae), with potential climate change

Insects in temperate regions are predicted to be at low risk of climate change relative to tropical species. However, these assumptions have generally been poorly examined in all regions, and such forecasting fails to account for microclimatic variation and behavioural optimisation. Here, we test how a population of the dominant ant species, Iridomyrmex purpureus, from temperate Australia responds to thermal stress. We show that ants regularly forage for short periods (minutes) at soil temperatures well above their upper thermal limits (upper lethal temperature = 45.8 ± 1.3 °C; CT_max = 46.1 °C) determined over slightly longer periods (hours) and do not show any signs of a classic thermal performance curve in voluntary locomotion across soil surface temperatures of 18.6–57°C (equating to a body temperature of 24.5–43.1 °C). Although ants were present all year round, and dynamically altered several aspects of their thermal biology to cope with low temperatures and seasonal variation, temperature-dependence of running speed remained invariant and ants were unable to elevate high temperature tolerance using plastic responses. Measurements of microclimate temperature were higher than ant body temperatures during the hottest part of the day, but exhibited a stronger relationship with each other than air temperatures from the closest weather station. Generally close associations of ant activity and performance with microclimatic conditions, possibly to maximise foraging times, suggest I. purpureus displays highly opportunistic thermal responses and readily adjusts behaviour to cope with high trail temperatures. Increasing frequency or duration of high temperatures is therefore likely to result in an immediate reduction in foraging efficiency. In summary, these results suggest that (1) soil-dwelling temperate insect populations may be at higher risks of thermal stress with increased frequency or duration of high temperatures resulting from climate change than previously thought, however, behavioural cues may be able to compensate to some extent; and (2) indices of climate change-related thermal stress, warming tolerance and thermal safety margin, are strongly influenced by the scale of climate metrics employed.     

Mutualism fails when climate response differs between interacting species

Successful species interactions require that both partners share a similar cue. For many species, spring warming acts as a shared signal to synchronize mutualist behaviors. Spring flowering plants and the ants that disperse their seeds respond to warming temperatures so that ants forage when plants drop seeds. However, where warm‐adapted ants replace cold‐adapted ants, changes in this timing might leave early seeds stranded without a disperser. We investigate plant seed dispersal south and north of a distinct boundary between warm‐ and cold‐adapted ants to determine if changes in the ant species influence local plant dispersal. The warm‐adapted ants forage much later than the cold‐adapted ants, and so we first assess natural populations of early and late blooming plants. We then transplant these plants south and north of the ant boundary to test whether distinct ant climate requirements disrupt the ant–plant mutualism. Whereas the early blooming plant's inability to synchronize with the warm‐adapted ant leaves its populations clumped and patchy and its seedlings clustered around the parents in natural populations, when transplanted into the range of the cold‐adapted ant, effective seed dispersal recovers. In contrast, the mutualism persists for the later blooming plant regardless of location because it sets seed later in spring when both warm‐ and cold‐adapted ant species forage, resulting in effective seed dispersal. These results indicate that the climate response of species interactions, not just the species themselves, is integral in understanding ecological responses to a changing climate. Data linking phenological synchrony and dispersal are rare, and these results suggest a viable mechanism by which a species' range is limited more by biotic than abiotic interactions – despite the general assumption that biotic influences are buried within larger climate drivers. These results show that biotic partner can be as fundamental a niche requirement as abiotic resources.     

Global variation in thermal tolerances and vulnerability of endotherms to climate change

The relationships among species'' physiological capacities and the geographical variation of ambient climate are of key importance to understanding the distribution of life on the Earth. Furthermore, predictions of how species will respond to climate change will profit from the explicit consideration of their physiological tolerances. The climatic variability hypothesis, which predicts that climatic tolerances are broader in more variable climates, provides an analytical framework for studying these relationships between physiology and biogeography. However, direct empirical support for the hypothesis is mostly lacking for endotherms, and few studies have tried to integrate physiological data into assessments of species'' climatic vulnerability at the global scale. Here, we test the climatic variability hypothesis for endotherms, with a comprehensive dataset on thermal tolerances derived from physiological experiments, and use these data to assess the vulnerability of species to projected climate change. We find the expected relationship between thermal tolerance and ambient climatic variability in birds, but not in mammals—a contrast possibly resulting from different adaptation strategies to ambient climate via behaviour, morphology or physiology. We show that currently most of the species are experiencing ambient temperatures well within their tolerance limits and that in the future many species may be able to tolerate projected temperature increases across significant proportions of their distributions. However, our findings also underline the high vulnerability of tropical regions to changes in temperature and other threats of anthropogenic global changes. Our study demonstrates that a better understanding of the interplay among species'' physiology and the geography of climate change will advance assessments of species'' vulnerability to climate change.     

Characterization of the thermal tolerances of forest ants of New England

Characterization of thermal tolerances of ants, which are both abundant and important in most terrestrial ecosystems, is needed since thermal constraints can inform how a species may respond to local climatic change. Here we identified the thermal tolerances of 16 common ant species of the Northeastern United States and determined relationships between body size, desiccation, and thermal tolerance among species. We hypothesized that maximum heat tolerances of these species would differ and be related to body size and capacity to resist desiccation. We identified four distinct groups of species belonging to one of three subfamilies, Dolichoderinae, Formicinae, or Myrmicinae, with different maximum thermal tolerances. Group “a” had a mean thermal tolerance of approximately 43°C (±1°C), group “b” had a mean thermal tolerance of 40°C (±1°C), group “c” had a mean thermal tolerance of 38°C (±0°C), and group “d” had a mean thermal tolerance of 36°C (±0°C). Groups “a” and “d” consisted of a single species (in the subfamilies Myrmicinae and Formicinae, respectively), while groups “b” and “c” were a mix of species in the subfamilies Myrmicinae, Formicinae, and Dolichoderinae. In the subfamily Formicinae, thermal tolerance increased with body size and critical water content, a metric of desiccation tolerance. In contrast, in the subfamily Myrmicinae, higher thermal tolerance was correlated with intermediate body size and lower critical water content. These findings suggest that the two dominant subfamilies in Northeastern deciduous forests have different relationships between body size, capacity to tolerate desiccation, and thermal tolerances across species. This variation in thermal tolerance suggests that climatic change may impact species differently.     

Oil palm expansion into rain forest greatly reduces ant biodiversity in canopy,epiphytes and leaf-litter

Oil palm cultivation is expanding rapidly into many of the world's most biodiverse tropical regions. One of the most functionally important and ecologically dominant animal groups in these environments is the ants. Here, we quantify the overall impacts of clear-felling lowland dipterocarp rainforest and conversion into oil palm plantation on ant diversity. At study sites in Sabah, Malaysia we collected ants from three microhabitats: 1 – the canopy, 2 – bird's nest ferns (Asplenium nidus complex, a common epiphyte in forest and oil palm), and 3 – leaf litter. We also measured temperature, humidity and light at collection sites to assess their impacts on ant community composition. Total ant species richness decreased from 309 to 110 (?64%) between forest and oil palm plantation. However, this impact was not the same across all microhabitats, with bird's nest ferns maintaining almost the same number of ant species in oil palm compared to forest (forest-oil palm, ferns: 36–35 (3% loss), canopy: 120–58 (52% loss), leaf litter: 216–56 (74% loss)). Relative abundance distributions remained the same for fern-dwelling ants, but became less even for oil palm ants in both the canopy and the leaf litter. These differences may be due in part to the ability of bird's nest ferns to provide a stable microclimate in hot, dry plantations. We also found that non-native ant species were more abundant in oil palm than in forest, and few forest ant species survived in plantations in any of the microhabitats. Only 59 of the 309 forest species persisted in oil palm plantations, corresponding to an 81% loss of forest species resulting from habitat conversion. Although oil palm supports many more ant species than has been previously reported, converting forest into plantation still leads to a dramatic reduction in species richness. The maintenance of forested areas is therefore vital for the conservation of ant biodiversity.     

Knitting patterns of biodiversity,range size and body size in aquatic beetle faunas: significant relationships but slightly divergent drivers

1. Ecogeographical rules refer to recurring patterns in nature, including the latitudinal diversity gradient (LDG), Rapoport's rule and Bergmann's rule, amongst others. In the present study, the existence of these rules was examined for diving beetles (Coleoptera: Dytiscidae), a family of aquatic predatory beetles. 2. Assemblage‐level data were analysed for diving beetles, focusing on species richness, local contribution to beta diversity (LCBD), mean range size and mean body size across the biogeographical provinces of Northern Europe. First, each of these variables was correlated with latitude, and then variation in each variable was modelled using actual environmental variables in boosted regression tree analysis. 3. Species richness was found to decrease with latitude, LCBD increased with latitude, mean range size did not show a significant relationship with latitude, and mean body size decreased with latitude. The latter finding was in contrast to Bergmann's rule. The actual environmental variables best predicting variation in these four response variables varied among the models, although they generally included temperature‐related and land use variables as the most influential ones. 4. The results obtained in the present study suggest that diving beetles conformed to the LDG, did not follow Rapoport's rule, and showed a reversed latitudinal gradient in the context of Bergmann's rule. In addition, species‐poor provinces harboured ecologically most unique faunas, suggesting that species richness and LCBD are complementary measures of biodiversity. 5. Even though general support was not found for most of the ecogeographical rules examined, the findings of the present study are interesting because they suggest that aquatic ectothermic invertebrates may show patterns different from those originally described for terrestrial endothermic vertebrates.     

Caterpillars escape predation in habitat and thermal refuges

1. Climate and, therefore, abiotic conditions, are changing rapidly, and many ecological interactions depend on them. In this study, how abiotic conditions mediate a predator–prey interaction were examined. 2. Caterpillars of Platyprepia virginalis (Boisduval) (Arctiidae) were found previously to be more abundant in wet habitats and thick litter cover compared with drier habitats and little or no litter. We hypothesised that wet litter provided caterpillars with refuges from an important ant predator, Formica lasioides. It was further hypothesised that caterpillars would be able to move at lower temperatures than ants, thus providing them with a thermal refuge. 3. In the lab, caterpillars were more likely to escape ant predation and survive on wet litter and at lower temperatures. At all temperatures, ant recruitment was lower in wet litter than dry litter although ants were more active on litter than bare soil. Thus, wet litter may serve as a habitat refuge for caterpillars from ants. 4. Caterpillars were able to maintain activity at temperatures 8–14 °C lower than F. lasioides. Thus colder temperatures may serve as a thermal refuge for caterpillars from ants. 5. It was hypothesised that caterpillars can escape ant predation when precipitation causes wet litter and at temperatures that they experience commonly in the field. This mismatch between caterpillars and their predators in ability to tolerate wet litter and low temperatures may affect their field distribution and abundance. Expected future warmer and drier conditions may not provide these refuges.     

Niche breadth,environmental landscape,and physical barriers: their importance as determinants of species distributions

Species distributions and their patterns in geographical space have been studied for several decades and explained by theories such as Janzen's, with respect to the nature of dispersal barriers in the Tropics, and Rapoport's, with respect to range size. However, the roles of specific environmental and geographical factors (e.g. ecological niche breadth, geographical barriers, etc.) in shaping species ranges and distributional patterns remain largely unexplored. The present study analyzed predictions from these two theories via analysis of virtual species with respect to biogeographical patterns: virtual species were created across South America, covering all major environments on the continent, and were used to compare effects of niche breadth, environmental availability, connectivity, seasonality, and the presence of known biogeographical barriers (rivers) in shaping species distributions and biodiversity patterns. Geographical ranges varied from narrow to broad, depending on the location of the seed point when comparing species produced with the same niche breadth. Analysis without consideration of seasonality and barriers produced species with broader distributions in the Tropics and narrower distributions in montane and temperate regions of the continent. When seasonality was included, however, broader ranges were concentrated in temperate regions, thus supporting Janzen's idea. Rapoport's rule of broader geographical ranges at higher latitudes was supported only when seasonality and physical barriers were included but not in species with very narrow or very broad niches, suggesting that this ‘rule’ results from interactions among niche breadth, dispersal capabilities, and dispersal barriers. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 241–250.     

Size–abundance relationships in Florida ant communities reveal how ants break the energetic equivalence rule

1. Ants are among the most abundant terrestrial organisms, yet little is known of how ant communities divide resources because it is difficult to measure the number of individuals in colonies and the density of colonies. 2. The body size–abundance relationships of the ants of five upland ecosystems in Florida were examined. The study tested whether abundance, energy use, and total biomass were distributed among species and body sizes as predicted by Damuth's energetic equivalence rule. Estimates of average worker body size, colony size, colony mass, and field metabolic rates were used to examine the relationships among body sizes, energy use, and total biomass. 3. Analyses revealed significant variation in energy use and did not support the energetic equivalence hypothesis. Specifically, the energy use and total standing biomass of species with large workers and colonies was much greater than smaller species. 4. These results suggest that larger species with larger colonies account for a disproportionate fraction of the total abundance and biomass of ants. A general model of resource allocation in colonies provides a possible explanation for why ants do not conform to the predictions of the energetic equivalence rule and for why ants are so abundant.     

Linking species thermal tolerance to elevational range shifts in upland dung beetles

Climate warming has been proposed as the main cause of the recent range shifts seen in many species. Although species' thermal tolerances are thought to play a key role in determining responses to climate change, especially in ectotherms, empirical evidence is still limited. We investigate the connection between species' thermal tolerances, elevational range and shifts in the lower elevational limit of dung beetle species (Coleoptera, Aphodiidea) in an upland region in the northwest of England. We measured thermal tolerances in the laboratory, and used current and historical distribution data to test specific hypotheses about the area's three dominant species, particularly the species most likely to suffer from warming: Agollinus lapponum. We found marked differences between species in their minimum and maximum thermal tolerance and in their elevational range and patterns of abundance. Overall, differences in thermal limits among species matched the abundance patterns along the elevation gradient expected if distributions were constrained by climate. Agollinus lapponum abundance increased with elevation and this species showed lower maximum and minimum thermal limits than Acrossus depressus, for which abundance declined with elevation. Consistent with lower tolerance to high temperature, we recorded an uphill retreat of the low elevation limit of A. lapponum (177 m over 57 yr) in line with the increase in summer temperature observed in the region over the same period. Moreover, this species has been replaced at low and mid‐elevations by the other two warm‐tolerant species (A. depressus and Agrilinus ater). Our results provide empirical evidence that species' thermal tolerance constrains elevational ranges and contributes to explain the observed responses to climate warming. A mechanistic understanding of how climate change directly affects species, such as the one presented here, will provide a robust base to inform predictions of how individual species and whole assemblages may change in the future.