Environmental context alters ecological trade‐offs controlling ant coexistence in a spatially heterogeneous region

1. Ecological trade‐offs in ant (Hymenoptera: Formicidae) assemblages and their implications for coexistence boast a rich history in entomology. Yet investigations of trade‐offs have largely been limited to homogeneous environments. We examined how environmental context modifies trade‐off expression in an ant assemblage spanning a heterogeneous region in central Florida, U.S.A. 2. We examined how trade‐off expression is altered among two contrasting habitat types: open shrub and forest. We tested for the presence of the dominance‐discovery trade‐off and two dominance‐thermal tolerance trade‐offs by estimating behavioral dominance, discovery ability, and thermal tolerance (foraging thermal limit, lethal temperature, and maximal abundance temperature) for a wide range of interacting ant species. 3. We found significantly linear dominance hierarchies in both shrub and forest habitats, showing dominant species out‐compete subordinates for food resources. In thermally stressful shrub habitats, subordinates exhibit higher thermal tolerances, take greater thermal risks, and reach maximum forager abundances at higher temperatures than do dominant species. This suggests temperature mediated trade‐offs control coexistence in shrub habitat. In thermally moderate forest habitat, we found limited evidence for trade‐offs between competitive dominance and resource discovery or between dominance and thermal traits, implying other processes control coexistence. These results demonstrate that trade‐offs controlling ant coexistence may be contingent on environmental context.     

Ecomorphology of plesiosaur flipper geometry

The Plesiosauria is an extinct group of marine reptiles once common in mesozoic seas. Previous work on plesiosaur hunting styles has suggested that short‐necked, large‐headed animals were pursuit predators, whereas long‐necked, small‐headed animals were ambush predators. This study presents new data on the aspect ratios (ARs) of plesiosaur flippers, and interprets these data via comparison with AR in birds, bats and aircraft. Performance trade‐offs implicit in AR variation are well‐understood in the context of aircraft design, and these trade‐offs have direct ecomorphological analogues in birds and bats. Knowledge of these trade‐offs allows interpretation of variation in plesiosaur AR. By analogy, short‐necked taxa were specialized for manoeuvrability and pursuit, whereas long‐necked taxa were generally specialized for efficiency and cruising. These interpretations agree with previous assessments of maximum swimming speed.     

Seasonality directs contrasting food collection behavior and nutrient regulation strategies in ants

Long-lived animals, including social insects, often display seasonal shifts in foraging behavior. Foraging is ultimately a nutrient consumption exercise, but the effect of seasonality per se on changes in foraging behavior, particularly as it relates to nutrient regulation, is poorly understood. Here, we show that field-collected fire ant colonies, returned to the laboratory and maintained under identical photoperiod, temperature, and humidity regimes, and presented with experimental foods that had different protein (p) to carbohydrate (c) ratios, practice summer- and fall-specific foraging behaviors with respect to protein-carbohydrate regulation. Summer colonies increased the amount of food collected as the p:c ratio of their food became increasingly imbalanced, but fall colonies collected similar amounts of food regardless of the p:c ratio of their food. Choice experiments revealed that feeding was non-random, and that both fall and summer ants preferred carbohydrate-biased food. However, ants rarely ate all the food they collected, and their cached or discarded food always contained little carbohydrate relative to protein. From a nutrient regulation strategy, ants consumed most of the carbohydrate they collected, but regulated protein consumption to a similar level, regardless of season. We suggest that varied seasonal food collection behaviors and nutrient regulation strategies may be an adaptation that allows long-lived animals to meet current and future nutrient demands when nutrient-rich foods are abundant (e.g. spring and summer), and to conserve energy and be metabolically more efficient when nutritionally balanced foods are less abundant.     

Trade‐offs in the evolution of bumblebee colony and body size: a comparative analysis

Trade‐offs between life‐history traits – such as fecundity and survival – have been demonstrated in several studies. In eusocial insects, the number of organisms and their body sizes can affect the fitness of the colony. Large‐than‐average body sizes as well as more individuals can improve a colony's thermoregulation, foraging efficiency, and fecundity. However, in bumblebees, large colonies and large body sizes depend largely on high temperatures and a large amount of food resources. Bumblebee taxa can be found in temperate and tropical regions of the world and differ markedly in their colony sizes and body sizes. Variation in colony size and body size may be explained by the costs and benefits associated with the evolutionary history of each species in a particular environment. In this study, we explored the effect of temperature and precipitation (the latter was used as an indirect indicator of food availability) on the colony and body size of twenty‐one bumblebee taxa. A comparative analysis controlling for phylogenetic effects as well as for the body size of queens, workers, and males in bumblebee taxa from temperate and tropical regions indicated that both temperature and precipitation affect colony and body size. We found a negative association between colony size and the rainiest trimester, and a positive association between the colony size and the warmest month of the year. In addition, male bumblebees tend to evolve larger body sizes in places where the rain occurs mostly in the summer and the overall temperature is warmer. Moreover, we found a negative relationship between colony size and body sizes of queens, workers, and males, suggesting potential trade‐offs in the evolution of bumblebee colony and body size.     

QUANTITATIVE GENETIC DIVERGENCE AND STANDING GENETIC (CO)VARIANCE IN THERMAL REACTION NORMS ALONG LATITUDE

Although the potential to adapt to warmer climate is constrained by genetic trade‐offs, our understanding of how selection and mutation shape genetic (co)variances in thermal reaction norms is poor. Using 71 isofemale lines of the fly Sepsis punctum, originating from northern, central, and southern European climates, we tested for divergence in juvenile development rate across latitude at five experimental temperatures. To investigate effects of evolutionary history in different climates on standing genetic variation in reaction norms, we further compared genetic (co)variances between regions. Flies were reared on either high or low food resources to explore the role of energy acquisition in determining genetic trade‐offs between different temperatures. Although the latter had only weak effects on the strength and sign of genetic correlations, genetic architecture differed significantly between climatic regions, implying that evolution of reaction norms proceeds via different trajectories at high latitude versus low latitude in this system. Accordingly, regional genetic architecture was correlated to region‐specific differentiation. Moreover, hot development temperatures were associated with low genetic variance and stronger genetic correlations compared to cooler temperatures. We discuss the evolutionary potential of thermal reaction norms in light of their underlying genetic architectures, evolutionary histories, and the materialization of trade‐offs in natural environments.     

Selection of food patches by sympatric herbivores in response to concealment and distance from a refuge

Small herbivores face risks of predation while foraging and are often forced to trade off food quality for safety. Life history, behaviour, and habitat of predator and prey can influence these trade‐offs. We compared how two sympatric rabbits (pygmy rabbit, Brachylagus idahoensis; mountain cottontail, Sylvilagus nuttallii) that differ in size, use of burrows, and habitat specialization in the sagebrush‐steppe of western North America respond to amount and orientation of concealment cover and proximity to burrow refuges when selecting food patches. We predicted that both rabbit species would prefer food patches that offered greater concealment and food patches that were closer to burrow refuges. However, because pygmy rabbits are small, obligate burrowers that are restricted to sagebrush habitats, we predicted that they would show stronger preferences for greater cover, orientation of concealment, and patches closer to burrow refuges. We offered two food patches to individuals of each species during three experiments that either varied in the amount of concealment cover, orientation of concealment cover, or distance from a burrow refuge. Both species preferred food patches that offered greater concealment, but pygmy rabbits generally preferred terrestrial and mountain cottontails preferred aerial concealment. Only pygmy rabbits preferred food patches closer to their burrow refuge. Different responses to concealment and proximity to burrow refuges by the two species likely reflect differences in perceived predation risks. Because terrestrial predators are able to dig for prey in burrows, animals like pygmy rabbits that rely on burrow refuges might select food patches based more on terrestrial concealment. In contrast, larger habitat generalists that do not rely on burrow refuges, like mountain cottontails, might trade off terrestrial concealment for visibility to detect approaching terrestrial predators. This study suggests that body size and evolutionary adaptations for using habitat, even in closely related species, might influence anti‐predator behaviors in prey species.     

Wildlife Recovery During Tropical Forest Succession: Assessing Ecological Drivers of Community Change

Despite the high proportion of secondary forests in the tropics, their conservation value remains poorly understood, particularly with regard to animals. Most theoretical studies of succession have focused on plants, linking life history trade‐offs to well‐known patterns of community change. However, the same trade‐offs proposed for plants should apply to animals, and indeed, animal studies show a change in community dominance from habitat generalist to forest specialist species during succession. Focusing on the diverse terrestrial small mammals of the endangered Atlantic Forest, we assessed which ecological drivers (habitat structure and food availability) affect community changes during succession. If the change in community dominance is driven by trade‐offs between productivity and efficiency, it should be mainly associated with a decrease in food availability. As expected, from younger to older forest, habitat generalists decreased in richness and total abundance, concurrent with a decrease in arthropod biomass. By contrast, the increase in richness and total abundance of forest specialists was not clearly supported by the data; however, this group was not affected by food availability. These results are congruent with a trade‐off between competitive ability and ability to use abundant resources, and indicate that the major community change during succession involves habitat generalists. Secondary forests may thus be valuable for conservation, at least where habitat loss and fragmentation are not high, and old growth forest is available.     

Anthropogenic food provisioning and immune phenotype: Association among supplemental food,body condition,and immunological parameters in urban environments

Direct or indirect supplemental feeding of free‐ranging animals occurs worldwide, resulting in significant impacts on population density or altered demographic processes. Another potential impact of increased energy intake from supplemental feeding is altered immunocompetence. As immune system maintenance is energetically costly, there may be trade‐offs between immune responses and other energy‐demanding physiological processes in individual animals. Although increased availability of food sources through supplemental feeding is expected to increase the overall immunocompetence of animals, empirical data verifying the association between supplemental feeding and different immune parameters are lacking. Understanding the potential influence of supplemental feeding on immune phenotypes is critical, as it may also impact host–pathogen dynamics in free‐ranging animals. Using urban stray cats as a study model, we tested for associations between the intensity of supplemental feeding due to cat caretaker activity (CCA); body condition; and immune phenotype (bacterial killing assay (BKA), immunoglobulin G (IgG) concentration, and leukocyte counts). Significantly higher bacterial killing ability was observed in cats from high CCA districts, whereas higher IgG concentration and eosinophil counts were observed in cats from low CCA districts. Other leukocyte counts and body condition indices showed no significant association with CCA. We observed varying patterns of different immune components in relation to supplemental feeding. Out data suggest that supplemental feeding influences immune phenotype, not only by means of energy provisioning, but also by potentially reducing exposure rates to parasite infections through stray cat behavioral changes.     

Can ant‐eating Zodarion spiders (Araneae: Zodariidae) develop on a diet optimal for euryphagous arthropod predators?

Abstract Little attention is paid to the behavioural and physiological adaptations of ant‐eating predators. It is expected that there should be a strong selection for traits related to prey handling, leading to the evolution of morphological, behavioural and nutritional adaptations. Such adaptations may then entail trade‐offs in handling and utilization of alternative prey. To investigate behavioural as well as nutritional adaptations and the occurrence of the corresponding trade‐offs in two ant‐eating spiders of the genus Zodarion [Zodarion atlanticum Pekár & Cardoso and Zodarion germanicum (C. L. Koch)], spiders are reared on two diets: ants (i.e. their preferred prey) and fruit flies (i.e. an alternative prey that is nutritionally optimal for euryphagous spiders). Food consumption is observed and several fitness‐related life‐history parameters are measured. Although spiders readily accept ants, more than one‐third of 35 spiders refuse to consume fruit flies and starve. Furthermore, severe hunger does not induce these individuals to accept fruit flies. Starving spiders die before moulting to the second stadium. Spiders that eat fruit flies increase only little and slowly in weight, and all of these die during the first two stadia. By contrast, spiders on an ant diet increase dramatically in weight, and develop up to the fourth stadium. These data indicate that fruit flies are not suitable for Zodarion, supporting the hypothesis that there are behavioural and nutritional trade‐offs. Taking into account the results of previous studies, it is suggested that nutritional trade‐offs are generally important for stenophagous spiders.     

Differences in food preferences between individuals and populations of domestic cats Felis silvestris catus

The food preferences of 64 cats, of which 28 were domestic pets and 36 were free-ranging animals on three farms, were tested using five food types; a sixth type was also tested on the farm cats. Information was also gathered on the background diet of the house cats (individually) and of each farm cat colony. Consistent differences were found between the predefined groups of cats in their preferences for three foods, Hard Dry (HD), Canned Meat (CM) and Raw Beef (RB), the greatest differences being between house cats and farm cats as a whole. The three colonies of farm cats showed distinct differences in preference, for RB and/or CM, and for the sixth food, Soft Dry (SD). Differences between farms could be explained by a reduced preference for items similar to major components of the background diet of each colony, i.e. selection in favour of foods that were temporally rare. Differences between house and farm cats, and within the house cats, could not be explained in this way; the house cats were neophobic towards RB, and the farm cats ate little of the HD food, possibly because they found it difficult to ingest.     

THE CONTRIBUTION OF SPONTANEOUS MUTATIONS TO THERMAL SENSITIVITY CURVE VARIATION IN DROSOPHILA SERRATA

Many traits studied in ecology and evolutionary biology change their expression in response to a continuously varying environmental factor. One well‐studied example are thermal performance curves (TPCs); continuous reaction norms that describe the relationship between organismal performance and temperature and are useful for understanding the trade‐offs involved in thermal adaptation. We characterized curves describing the thermal sensitivity of voluntary locomotor activity in a set of 66 spontaneous mutation accumulation lines in the fly Drosophila serrata. Factor‐analytic modeling of the mutational variance–covariance matrix, M , revealed support for three axes of mutational variation in males and two in females. These independent axes of mutational variance corresponded well to the major axes of TPC variation required for different types of thermal adaptation; “faster‐slower” representing changes in performance largely independent of temperature, and the “hotter‐colder” and “generalist‐specialist” axes, representing trade‐offs. In contrast to its near‐absence from standing variance in this species, a “faster‐slower” axis, accounted for most mutational variance (75% in males and 66% in females) suggesting selection may easily fix or remove these types of mutations in outbred populations. Axes resembling the “hotter‐colder” and “generalist‐specialist” modes of variation contributed less mutational variance but nonetheless point to an appreciable input of new mutations that may contribute to thermal adaptation.     

Ant assemblages on littered food waste and food removal rates in urban–suburban parks of Tokyo

Ants are one of the major animals utilizing waste in urban areas, and presumably have an important role in nutrient redistribution and cycling. However, information on ant assemblages on artificial food and their food removal rates in different habitat types is lacking, and the relationship between ant assemblage and removal rate is poorly understood. We assessed assemblages of ants on experimentally placed foods (potato chips, cookies and ham) and their food removal rates on three land cover types (woodlands, lawns and pavements) at 90 sites in 10 urban and suburban parks of Tokyo. Then, we examined the relationship between ant assemblage and food removal rate. In total, 11 trophic generalist ants were associated with food removal. Species composition differed significantly between land cover types, and the mean number of species was higher in woodlands than on lawns and pavements, while not significantly different between urban and suburban parks. The ants removed, on average, 6.3 g of 18-g food in fresh weight (or 5.3 g of 13.9-g food in dry weight) per site in 24 h. There were both negative and positive associations between ant species presence and food removal rate, but no significant associations between species number and food removal rate. Food removal rate was highest on lawns, and that in woodlands was not significantly different from that on lawns and pavements. Therefore, our study suggests that ants on lawns and pavements, despite lower number of species and altered species composition, have a comparable or higher ability to redistribute nutrients from artificial foods as compared with woodland ants. This indicates that such highly artificial land cover types should also be included in studies on urban ecosystem services.     

Temperature,size, reproductive allocation,and life‐history evolution in a gregarious caterpillar

The present study aimed to investigate the relationship between growth rate, final mass, and larval development, as well as how this relationship influences reproductive trade‐offs, in the context of a gregarious life‐style and the need to keep an optimal group size. We use as a model two sympatric populations of the pine processionary moth Thaumetopoea pityocampa, which occur in different seasons and thus experience different climatic conditions. Thaumetopoea pityocampa is a strictly gregarious caterpillar throughout the larval period, which occurs during winter in countries all over the Mediterranean Basin. However, in 1997, a population in which larval development occurs during the summer was discovered in Portugal, namely the summer population (SP), as opposed to the normal winter population (WP), which coexists in the same forest feeding on the same host during the winter. Both populations were monitored over 3 years, with an assessment of the length of the larval period and its relationship with different climatic variables, final mass and adult size, egg size and number, colony size, and mortality at different life stages. The SP larval period was reduced as a result of development in the warmer part of the year, although it reached the same final mass and adult size as the WP. Despite an equal size at maturity, a trade‐off between egg size and number was found between the two populations: SP produced less but bigger eggs than WP. This contrasts with the findings obtained in other Lepidoptera species, where development in colder environments leads to larger eggs at the expense of fecundity, but corroborates the trend found at a macro‐geographical scale for T. pityocampa, with females from northern latitudes and a colder environment producing more (and smaller) eggs. The results demonstrate the importance of the number of eggs in cold environments as a result of an advantage of large colonies when gregarious caterpillars develop in such environments, and these findings are discussed in accordance with the major theories regarding size in animals. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 340–349.     

The flow of food in colonies of the fire ant, Solenopsis invicta: a multifactorial study

ABSTRACT. A study was conducted to determine the interaction of several factors in the trophallactic distribution of food in colonies of Solenopsis invicta Buren. Radioactive iodine (¹²⁵I) was used to quantify both the ingestion of test foods by foragers and the distribution of this food to nestmates. A multifactorial design tested the simultaneous effects of: (1) food-type (sugar water, casein hydrolysate, or oil), (2) temperature during foraging and food-sharing (25, 30 or 35°C), and (3) starvation period (colonies deprived for 0, 3, 7 or 14 days). The amount of food consumed by foragers was influenced by the interaction of food-type and starvation. Foragers from severely starved colonies ingested more aqueous foods than did those from slightly starved colonies, but oil was always taken in smaller quantities and its ingestion was least affected by starvation. Ingestion was generally lowest at 35°C, probably because of mutual interference of workers while feeding. Within the colony, food sharing varied with food type and increased with starvation. The sugar solution was utilized primarily by workers. Amino acids were shared throughout the colony, but were directed preferentially to the growing larvae. Soy oil was equally shared among workers and larvae and reached more colony members per initial unit volume than did the other foods. The test factors also interacted in their influence on trophallaxis. Starvation stimulated the dispersal of labelled foods, but at different intensities for different foods. The probability of the queen's receiving food increased when the labelled food was widely exchanged among her nestmates. Casein hydrolysate reached the queen in more cases than did the other tested foods.     

Mice prefer rare food

In the first experiment designed to test directly for frequency-dependent selection of food by a mammal, mice were presented with pairs of food in 9: 1 and 1:9 combinations. The foods used were crumbs of laboratory rat cake flavoured vanilla or peppermint and dyed green or brown: vanilla/green was presented with peppermint/brown, and vanilla/brown with peppermint/green. Vanilla was preferred to peppermint and green to brown: peppermint/brown was particularly unacceptable. Each type was preferred more when rare than when common. Preferences changed over the five days of the experiment, the preference for greens becoming less strong. Since the experimental trials were carried out in total darkness, the preferences must have been based on non-visual clues, indicating that the dyes were not odourless and tasteless to mice, as they are to man.     

Push and bite: trade‐offs between burrowing and biting in a burrowing skink (Acontias percivali)

Trade‐offs are thought to be important in constraining evolutionary divergence, as they may limit phenotypic diversification. Limbless animals that burrow head‐first have been suggested to be evolutionarily constrained in the development of a large head size and sexual head shape dimorphism because of potential trade‐offs associated with burrowing. Here we use an acontiine skink (Acontias percivali) to test for the existence of trade‐offs between traits thought to be important in burrowing (speed and force). As head size dimorphism has been shown to be limited in acontiine lizards, thus suggesting constraints on head size and shape, we additionally explore the potential for trade‐offs between burrowing and biting. Our data show that A. percivali uses a burrowing style different from those previously described for caecilians and amphisbaenians, which relies on the use of extensive lateral and dorsoventral head movements. Our data also show that animals use their entire bodies to generate force, as peak force was determined by total length only. Additionally, both bite force and the time needed to burrow into the substrate were principally determined by relative head width, suggesting a trade‐off between biting and burrow speed. Performance data were indeed suggestive of a correlation between bite force and the time needed to burrow, but additional data are needed to confirm this pattern. In summary, our data suggests that trade‐offs may exist, and may have been of crucial importance in shaping the evolution of head shape in A. percivali, and burrowing lizards more generally. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 91–99.     

Defensive traits exhibit an evolutionary trade‐off and drive diversification in ants

Evolutionary biologists have long predicted that evolutionary trade‐offs among traits should constrain morphological divergence and species diversification. However, this prediction has yet to be tested in a broad evolutionary context in many diverse clades, including ants. Here, we reconstruct an expanded ant phylogeny representing 82% of ant genera, compile a new family‐wide trait database, and conduct various trait‐based analyses to show that defensive traits in ants do exhibit an evolutionary trade‐off. In particular, the use of a functional sting negatively correlates with a suite of other defensive traits including spines, large eye size, and large colony size. Furthermore, we find that several of the defensive traits that trade off with a sting are also positively correlated with each other and drive increased diversification, further suggesting that these traits form a defensive suite. Our results support the hypothesis that trade‐offs in defensive traits significantly constrain trait evolution and influence species diversification in ants.     

Considering land–sea interactions and trade‐offs for food and biodiversity

With the human population expected to near 10 billion by 2050, and diets shifting towards greater per‐capita consumption of animal protein, meeting future food demands will place ever‐growing burdens on natural resources and those dependent on them. Solutions proposed to increase the sustainability of agriculture, aquaculture, and capture fisheries have typically approached development from single sector perspectives. Recent work highlights the importance of recognising links among food sectors, and the challenge cross‐sector dependencies create for sustainable food production. Yet without understanding the full suite of interactions between food systems on land and sea, development in one sector may result in unanticipated trade‐offs in another. We review the interactions between terrestrial and aquatic food systems. We show that most of the studied land–sea interactions fall into at least one of four categories: ecosystem connectivity, feed interdependencies, livelihood interactions, and climate feedback. Critically, these interactions modify nutrient flows, and the partitioning of natural resource use between land and sea, amid a backdrop of climate variability and change that reaches across all sectors. Addressing counter‐productive trade‐offs resulting from land‐sea links will require simultaneous improvements in food production and consumption efficiency, while creating more sustainable feed products for fish and livestock. Food security research and policy also needs to better integrate aquatic and terrestrial production to anticipate how cross‐sector interactions could transmit change across ecosystem and governance boundaries into the future.     

Temperature-mediated coexistence in temperate forest ant communities

Patterns of ant species diversity are well documented and yet the mechanisms promoting species coexistence among communities are often elusive. Two emerging hypotheses that account for coexistence in ant communities are the discovery-dominance tradeoff and the dominance-thermal tolerance tradeoff. Here we used behavioural assays and community-level sampling from ant assemblages in the southern Appalachians, USA to test for the discovery-dominance and dominance-thermal tolerance tradeoffs. Species that were behaviorally dominant during interspecific interactions tended to forage in a narrow window of generally warmer temperatures, whereas subordinate species tended to forage in a wide range of temperatures, including colder temperatures. Species that foraged at lower temperature tended to be behaviourally subordinate, suggesting that a dominance-thermal tolerance tradeoff promotes coexistence in this system. Species richness was positively related to site average annual temperature and within-site variation in ground temperature, suggesting that temperature also shapes the structure of ant communities and regulates diversity. There was no relationship between the ability of a species to discover food resources and its behavioural dominance, contrary to the predictions of the discovery-dominance tradeoff hypothesis. In sum, our results show that temperature plays numerous roles in promoting regional coexistence in this system.