We study male parentage and between-colony variation in sex allocation and sexual production in the desert ant Crematogaster smithi, which usually has only one singly-mated queen per nest. Colonies of this species are known to temporarily store nutrients in the large fat body of intermorphs, a specialized female caste intermediate in morphology between queens and workers. Intermorphs repackage at least part of this fat into consumable but viable male-destined eggs. If these eggs sometimes develop instead of being eaten, intermorphs will be reproductive competitors of the queen but—due to relatedness asymmetries—allies of their sister worker. Using genetic markers we found a considerable proportion of non-queen sons in some, but not all, colonies. Even though intermorphs produce ∼1.7× more eggs than workers, their share in the parentage of adult males is estimated to be negligible due to their small number compared to workers. Furthermore, neither colony-level sex allocation nor overall sexual production was correlated with intermorph occurrence or number. We conclude that intermorph-laid eggs typically do not survive and that the storage of nutrients and their redistribution as eggs by intermorphs is effectively altruistic.
Following its introduction from Asia to the USA, the Asian needle ant (Pachycondyla chinensis) is rapidly spreading into a wide range of habitats with great negative ecological affects. In addition, the species is a concern for human health because of its powerful, sometimes deadly, sting. Here, we assessed the potential of P. chinensis to spread further and to invade entirely new regions. We used species distribution models to assess suitable areas under current climatic conditions and in 2020, 2050 and 2080. With a consensus model, combining five different modelling techniques, three Global Circulation (climatic) Models and two CO2 emission scenarios, we generated world maps with suitable climatic conditions. Our models suggest that the species currently has a far greater potential distribution than its current exotic range, including large parts of the world landmass, including Northeast America, Southeast Asia and Southeast America. Climate change is predicted to greatly exacerbate the risk of P. chinensis invasion by increasing the suitable landmass by 64.9% worldwide, with large increases in Europe (+210.1%), Oceania (+75.1%), North America (+74.9%) and Asia (+62.7%). The results of our study suggest P. chinensis deserves increased attention, especially in the light of on-going climate change.
Environmental conditions and physical constraints both influence an animal's behavior. We investigate whether behavioral variation among colonies of the black harvester ant, Messor andrei, remains consistent across foraging and disturbance situations and ask whether consistent colony behavior is affected by nest site and weather. We examined variation among colonies in responsiveness to food baits and to disturbance, measured as a change in numbers of active ants, and in the speed with which colonies retrieved food and removed debris. Colonies differed consistently, across foraging and disturbance situations, in both responsiveness and speed. Increased activity in response to food was associated with a smaller decrease in response to alarm. Speed of retrieving food was correlated with speed of removing debris. In all colonies, speed was greater in dry conditions, reducing the amount of time ants spent outside the nest. While a colony occupied a certain nest site, its responsiveness was consistent in both foraging and disturbance situations, suggesting that nest structure influences colony personality.
behavioral syndromes; collective behavior; harvester ant; Messor andrei; nest structure; personality; plasticity; social insects; temperament
Argentine ants (Linepithema humile) live in groups of nests connected by trails to each other and to stable food sources. In a field study, we investigated whether some ants recruit directly from established, persistent trails to food sources, thus accelerating food collection. Our results indicate that Argentine ants recruit nestmates to food directly from persistent trails, and that the exponential increase in the arrival rate of ants at baits is faster than would be possible if recruited ants traveled from distant nests. Once ants find a new food source, they walk back and forth between the bait and sometimes share food by trophallaxis with nestmates on the trail. Recruiting ants from nearby persistent trails creates a dynamic circuit, like those found in other distributed systems, which facilitates a quick response to changes in available resources.
Collective behavior is produced by interactions among individuals. Differences among groups in individual response to interactions can lead to ecologically important variation among groups in collective behavior. Here we examine variation among colonies in the foraging behavior of the harvester ant, Pogonomyrmex barbatus. Previous work shows how colonies regulate foraging in response to food availability and desiccation costs: the rate at which outgoing foragers leave the nest depends on the rate at which foragers return with food. To examine how colonies vary in response to humidity and in foraging rate, we performed field experiments that manipulated forager return rate in 94 trials with 17 colonies over 3 years. We found that the effect of returning foragers on the rate of outgoing foragers increases with humidity. There are consistent differences among colonies in foraging activity that persist from year to year.
Urbanisation typically results in a reduction of hollow-bearing trees and an increase in the density of particularly species, potentially resulting in an increased level of competition as cavity-nesting species compete for a limited resource. To improve understanding of hollow usage between urban cavity-nesting species in Australia, particularly parrots, we investigated how the hollow-using assemblage, visitation rate, diversity and number of interactions varied between hollows within urban remnant forest and continuous forest. Motion-activated video cameras were installed, via roped access to the canopy, and hollow usage was monitored at 61 hollows over a two-year period. Tree hollows within urban remnants had a significantly different assemblage of visitors to those in continuous forest as well as a higher rate of visitation than hollows within continuous forest, with the rainbow lorikeet making significantly more visitations than any other taxa. Hollows within urban remnants were characterised by significantly higher usage rates and significantly more aggressive interactions than hollows within continuous forest, with parrots responsible for almost all interactions. Within urban remnants, high rates of hollow visitation and both interspecific and intraspecific interactions observed at tree hollows suggest the number of available optimal hollows may be limiting. Understanding the usage of urban remnant hollows by wildlife, as well as the role of parrots as a potential flagship for the conservation of tree-hollows, is vital to prevent a decrease in the diversity of urban fauna, particularly as other less competitive species risk being outcompeted by abundant native species.
Identifying mechanisms governing the establishment and spread of invasive species is a fundamental challenge in invasion biology. Because species invasions are frequently observed only after the species presents an environmental threat, research identifying the contributing agents to dispersal and subsequent spread are confined to retrograde observations. Here, we use a combination of seasonal surveys and experimental approaches to test the relative importance of behavioral and abiotic factors in determining the local co-occurrence of two invasive ant species, the established Argentine ant (Linepithema humile Mayr) and the newly invasive Asian needle ant (Pachycondyla chinensis Emery). We show that the broader climatic envelope of P. chinensis enables it to establish earlier in the year than L. humile. We also demonstrate that increased P. chinensis propagule pressure during periods of L. humile scarcity contributes to successful P. chinensis early season establishment. Furthermore, we show that, although L. humile is the numerically superior and behaviorally dominant species at baits, P. chinensis is currently displacing L. humile across the invaded landscape. By identifying the features promoting the displacement of one invasive ant by another we can better understand both early determinants in the invasion process and factors limiting colony expansion and survival.
Confirmation bias is a tendency of people to interpret information in a way that confirms their expectations. A long recognized phenomenon in human psychology, confirmation bias can distort the results of a study and thus reduce its reliability. While confirmation bias can be avoided by conducting studies blind to treatment groups, this practice is not always used. Surprisingly, this is true of research in animal behaviour, and the extent to which confirmation bias influences research outcomes in this field is rarely investigated. Here we conducted a meta-analysis, using studies on nestmate recognition in ants, to compare the outcomes of studies that were conducted blind with those that were not. Nestmate recognition studies typically perform intra- and inter colony aggression assays, with the a priori expectation that there should be little or no aggression among nestmates. Aggressive interactions between ants can include subtle behaviours such as mandible flaring and recoil, which can be hard to quantify, making these types of assays prone to confirmation bias. Our survey revealed that only 29% of our sample of 79 studies were conducted blind. These studies were more likely to report aggression among nestmates if they were conducted blind (73%) than if they were not (21%). Moreover, we found that the effect size between nestmate and non-nestmate treatment means is significantly lower in experiments conducted blind than those in which colony identity is known (1.38 versus 2.76). We discuss the implications of the impact of confirmation bias for research that attempts to obtain quantitative synthesises of data from different studies.
Social insect colonies operate without central control or any global assessment of what needs to be done by workers. Colony organization arises from the responses of individuals to local cues. Red harvester ants (Pogonomyrmex barbatus) regulate foraging using interactions between returning and outgoing foragers. The rate at which foragers return with seeds, a measure of food availability, sets the rate at which outgoing foragers leave the nest on foraging trips. We used mimics to test whether outgoing foragers inside the nest respond to the odor of food, oleic acid, the odor of the forager itself, cuticular hydrocarbons, or a combination of both with increased foraging activity. We compared foraging activity, the rate at which foragers passed a line on a trail, before and after the addition of mimics. The combination of both odors, those of food and of foragers, is required to stimulate foraging. The addition of blank mimics, mimics coated with food odor alone, or mimics coated with forager odor alone did not increase foraging activity. We compared the rates at which foragers inside the nest interacted with other ants, blank mimics, and mimics coated with a combination of food and forager odor. Foragers inside the nest interacted more with mimics coated with combined forager/seed odors than with blank mimics, and these interactions had the same effect as those with other foragers. Outgoing foragers inside the nest entrance are stimulated to leave the nest in search of food by interacting with foragers returning with seeds. By using the combined odors of forager cuticular hydrocarbons and of seeds, the colony captures precise information, on the timescale of seconds, about the current availability of food.
The foraging behavior of the arboreal turtle ant, Cephalotes goniodontus, was studied in the tropical dry forest of western Mexico. The ants collected mostly plant-derived food, including nectar and fluids collected from the edges of wounds on leaves, as well as caterpillar frass and lichen. Foraging trails are on small pieces of ephemeral vegetation, and persist in exactly the same place for 4–8 days, indicating that food sources may be used until they are depleted. The species is polydomous, occupying many nests which are abandoned cavities or ends of broken branches in dead wood. Foraging trails extend from trees with nests to trees with food sources. Observations of marked individuals show that each trail is travelled by a distinct group of foragers. This makes the entire foraging circuit more resilient if a path becomes impassable, since foraging in one trail can continue while a different group of ants forms a new trail. The colony’s trails move around the forest from month to month; from one year to the next, only one colony out of five was found in the same location. There is continual searching in the vicinity of trails: ants recruited to bait within 3 bifurcations of a main foraging trail within 4 hours. When bait was offered on one trail, to which ants recruited, foraging activity increased on a different trail, with no bait, connected to the same nest. This suggests that the allocation of foragers to different trails is regulated by interactions at the nest.
In Ensifera, the lack of well-supported phylogeny and the focus on acoustic communication of the terminal taxa hinders understanding of the evolutionary history of their signalling behaviour and the related sensory structures. For Rhaphidophoridae, the most relic of ensiferans following morphology-based phylogenies, the signalling modes are still unknown. Together with a detailed description of their mating process, we provide evidence on vibratory signalling for the sympatric European species Troglophilus neglectus and T. cavicola. Despite their temporal shift in reproduction, the species’ behaviours differ significantly. Signalling by abdominal vibration constitutes an obligatory part of courtship in T. neglectus, while it is absent in T. cavicola. Whole-body vibration is expressed after copulation in both species. While courtship signalling appears to stimulate females for mating, the function of post-copulation signals remains unclear. Mating and signalling of both species were found to take place in most cases on bark, and less frequently on other available substrates, like moss and rock. The signals’ frequency spectra were substrate dependent, but with the dominant peak always expressed below 120 Hz. On rock, the intensity of T. neglectus courtship signals was below the species’ physiological detection range, presumably constraining the evolution of such signalling in caves. The species’ behavioural divergence appears to reflect their divergent mating habitats, in and outside caves. We propose that short-range tremulation signalling in courtship, such as is expressed by T. neglectus, represents the primitive mode and context of mechanical signalling in Ensifera. The absence of high-frequency components in the signals may be related to the absence of the crista acoustica homologue (CAH) in the vibratory tibial organ of Rhaphidophoridae. This indirectly supports the hypothesis proposing that the CAH, as an evolutionary precursor of the ear, evolved in Ensifera along the (more) complex vibratory communication, also associated with signals of higher carrier frequency.
Many dynamical networks, such as the ones that produce the collective behavior of social insects, operate without any central control, instead arising from local interactions among individuals. A well-studied example is the formation of recruitment trails in ant colonies, but many ant species do not use pheromone trails. We present a model of the regulation of foraging by harvester ant (Pogonomyrmex barbatus) colonies. This species forages for scattered seeds that one ant can retrieve on its own, so there is no need for spatial information such as pheromone trails that lead ants to specific locations. Previous work shows that colony foraging activity, the rate at which ants go out to search individually for seeds, is regulated in response to current food availability throughout the colony's foraging area. Ants use the rate of brief antennal contacts inside the nest between foragers returning with food and outgoing foragers available to leave the nest on the next foraging trip. Here we present a feedback-based algorithm that captures the main features of data from field experiments in which the rate of returning foragers was manipulated. The algorithm draws on our finding that the distribution of intervals between successive ants returning to the nest is a Poisson process. We fitted the parameter that estimates the effect of each returning forager on the rate at which outgoing foragers leave the nest. We found that correlations between observed rates of returning foragers and simulated rates of outgoing foragers, using our model, were similar to those in the data. Our simple stochastic model shows how the regulation of ant colony foraging can operate without spatial information, describing a process at the level of individual ants that predicts the overall foraging activity of the colony.
Social insect colonies operate without any central control. Their collective behavior arises from local interactions among individuals. Here we present a simple stochastic model of the regulation of foraging by harvester ant (Pogonomyrmex barbatus) colonies, which forage for scattered seeds that one ant can retrieve on its own, so there is no need for pheromone trails to specific locations. Previous work shows that colony foraging activity is regulated in response to current food availability, using the rate of brief antennal contacts inside the nest between foragers returning with food and outgoing foragers. Our feedback-based algorithm estimates the effect of each returning forager on the rate at which foragers leave the nest. The model shows how the regulation of ant colony foraging can operate without spatial information, describing a process at the level of individual ants that predicts the overall foraging activity of the colony.
Numerous studies have examined the effect of urbanization on species richness and most studies implicate urbanization as the major cause of biodiversity loss. However, no study has identified an explicit connection between urbanization and biodiversity loss as the impact of urbanization is typically inferred indirectly by comparing species diversity along urban-rural gradients at a single time point. A different approach is to focus on the temporal rather than the spatial aspect and perform “before and after” studies where species diversity is cataloged over time in the same sites. The current study examined changes in ant abundance and diversity associated with the conversion of natural habitats into urban habitats. Ant abundance and diversity were tracked in forested sites that became urbanized through construction and were examined at 3 time points - before, during, and after construction. On average, 4.3±1.2 unique species were detected in undisturbed plots prior to construction. Ant diversity decreased to 0.7±0.8 species in plots undergoing construction and 1.5±1.1 species in plots 1 year after construction was completed. With regard to species richness, urbanization resulted in the permanent loss of 17 of the 20 species initially present in the study plots. Recovery was slow and only 3 species were present right after construction was completed and 4 species were present 1 year after construction was completed. The second objective examined ant fauna recovery in developed residential lots based on time since construction, neighboring habitat quality, pesticide inputs, and the presence of invasive ants. Ant diversity was positively correlated with factors that promoted ecological recovery and negatively correlated with factors that promoted ecological degradation. Taken together, these results address a critical gap in our knowledge by characterizing the short- and long-term the effects of urbanization on the loss of ant biodiversity.
This study investigates variation in collective behavior in a natural population of colonies of the harvester ant, Pogonomyrmex barbatus. Harvester ant colonies regulate foraging activity to adjust to current food availability; the rate at which inactive foragers leave the nest on the next trip depends on the rate at which successful foragers return with food. This study investigates differences among colonies in foraging activity and how these differences are associated with variation among colonies in the regulation of foraging. Colonies differ in the baseline rate at which patrollers leave the nest, without stimulation from returning ants. This baseline rate predicts a colony's foraging activity, suggesting there is a colony-specific activity level that influences how quickly any ant leaves the nest. When a colony's foraging activity is high, the colony is more likely to regulate foraging. Moreover, colonies differ in the propensity to adjust the rate of outgoing foragers to the rate of forager return. Naturally occurring variation in the regulation of foraging may lead to variation in colony survival and reproductive success.
behavioral reaction norm; behavioral syndrome; individual variation
The evolutionary success of ants and other social insects is considered to be intrinsically linked to division of labor among workers. The role of the brains of individual ants in generating division of labor, however, is poorly understood, as is the degree to which interspecific variation in worker social phenotypes is underscored by functional neurobiological differentiation. Here we demonstrate that dimorphic minor and major workers of different ages from three ecotypical species of the hyperdiverse ant genus Pheidole have distinct patterns of neuropil size variation. Brain subregions involved in sensory input (optic and antennal lobes), sensory integration, learning and memory (mushroom bodies), and motor functions (central body and subesophageal ganglion) vary significantly in relative size, reflecting differential investment in neuropils that likely regulate subcaste- and age-correlated task performance. Worker groups differ in brain size and display patterns of altered isometric and allometric subregion scaling that affect brain architecture independently of brain size variation. In particular, mushroom body size was positively correlated with task plasticity in the context of both age- and subcaste-related polyethism, providing strong, novel support that greater investment in this neuropil increases behavioral flexibility. Our findings reveal striking levels of developmental plasticity and evolutionary flexibility in Pheidole worker neuroanatomy, supporting the hypothesis that mosaic alterations of brain composition contribute to adaptive colony structure and interspecific variation in social organization.
The invasive Argentine ant (Linepithema humile) is established worldwide and displaces native ant species. In northern California, however, the native winter ant (Prenolepis imparis) persists in invaded areas. We found that in aggressive interactions between the two species, P. imparis employs a potent defensive secretion. Field observations were conducted at P. imparis nest sites both in the presence and absence of L. humile. These observations suggested and laboratory assays confirmed that P. imparis workers are more likely to secrete when outnumbered by L. humile. Workers of P. imparis were also more likely to secrete near their nest entrances than when foraging on trees. One-on-one laboratory trials showed that the P. imparis secretion is highly lethal to L. humile, causing 79% mortality. The nonpolar fraction of the secretion was chemically analyzed with gas chromatography/mass spectrometry, and found to be composed of long-chain and cyclic hydrocarbons. Chemical analysis of dissected P. imparis workers showed that the nonpolar fraction is derived from the Dufour's gland. Based on these conclusions, we hypothesize that this chemical defense may help P. imparis to resist displacement by L. humile.
Social insects exhibit coordinated behaviour without central control. Local interactions among individuals determine their behaviour and regulate the activity of the colony. Harvester ants are recruited for outside work, using networks of brief antennal contacts, in the nest chamber closest to the nest exit: the entrance chamber. Here, we combine empirical observations, image analysis and computer simulations to investigate the structure and function of the interaction network in the entrance chamber. Ant interactions were distributed heterogeneously in the chamber, with an interaction hot-spot at the entrance leading further into the nest. The distribution of the total interactions per ant followed a right-skewed distribution, indicating the presence of highly connected individuals. Numbers of ant encounters observed positively correlated with the duration of observation. Individuals varied in interaction frequency, even after accounting for the duration of observation. An ant's interaction frequency was explained by its path shape and location within the entrance chamber. Computer simulations demonstrate that variation among individuals in connectivity accelerates information flow to an extent equivalent to an increase in the total number of interactions. Individual variation in connectivity, arising from variation among ants in location and spatial behaviour, creates interaction centres, which may expedite information flow.
agent-based model; movement pattern; network analysis; Pogonomyrmex barbatus; spatial behaviour; weighted degree
‘Devil's gardens’ are nearly pure stands of the myrmecophyte, Duroia hirsuta, that occur in Amazonian rainforests. Devil's gardens are created by Myrmelachista schumanni ants, which nest in D. hirsuta trees and kill other plants using formic acid as an herbicide. Here, we show that this ant–plant mutualism has an associated cost; by making devil's gardens, M. schumanni increases herbivory on D. hirsuta. We measured standing leaf herbivory on D. hirsuta trees and found that they sustain higher herbivory inside than outside devil's gardens. We also measured the rate of herbivory on nursery-grown D. hirsuta saplings planted inside and outside devil's gardens in ant-exclusion and control treatments. We found that when we excluded ants, herbivory on D. hirsuta was higher inside than outside devil's gardens. These results suggest that devil's gardens are a concentrated resource for herbivores. Myrmelachista schumanni workers defend D. hirsuta against herbivores, but do not fully counterbalance the high herbivore pressure in devil's gardens. We suggest that high herbivory may limit the spread of devil's gardens, possibly explaining why devil's gardens do not overrun Amazonian rainforests.
ant–plant interactions; density dependence; diminishing returns; Janzen–Connell hypothesis; pure stands; resource concentration hypothesis
Various invasive ant species have negatively affected reproductive success in birds by disrupting nest site selection, incubation patterns, food supply, and by direct predation on nestlings. Impacts can be particularly severe when non-native ants colonize seabird nesting islands where thousands of birds may nest in high densities on the ground or in burrows or crevices. Here we report on the first documented effects of Myrmica rubra, the European fire ant, on the reproduction of birds in its non-native range. We documented herring gulls (Larus argentatus) on Appledore Island, Maine, engaging in more erratic incubation behaviors at nests infested by the ants. Newly-hatched chicks in some nests were swarmed by ants, leading to rapid chick death. Due to high overall rates of chick mortality, survival probabilities did not vary between nests with and without ant activity, however chick growth rates were slower at nests with ants than at ant-free nests. Ant infestation likely leads to longer-term fitness consequences because slower growth rates early in life may ultimately lead to lower post-fledging survival probabilities.
Many ants rely on both visual cues and self-generated chemical signals for navigation, but their relative importance varies across species and context. We evaluated the roles of both modalities during colony emigration by Temnothorax rugatulus. Colonies were induced to move from an old nest in the center of an arena to a new nest at the arena edge. In the midst of the emigration the arena floor was rotated 60°around the old nest entrance, thus displacing any substrate-bound odor cues while leaving visual cues unchanged. This manipulation had no effect on orientation, suggesting little influence of substrate cues on navigation. When this rotation was accompanied by the blocking of most visual cues, the ants became highly disoriented, suggesting that they did not fall back on substrate cues even when deprived of visual information. Finally, when the substrate was left in place but the visual surround was rotated, the ants' subsequent headings were strongly rotated in the same direction, showing a clear role for visual navigation. Combined with earlier studies, these results suggest that chemical signals deposited by Temnothorax ants serve more for marking of familiar territory than for orientation. The ants instead navigate visually, showing the importance of this modality even for species with small eyes and coarse visual acuity.
Global warming is characterized by not only an increase in the daily mean temperature, but also a diel asymmetric pattern. However, most of the current studies on climate change have only concerned with the mean values of the warming trend. Although many studies have been conducted concerning the responses of insects to climate change, studies that address the issue of diel asymmetric warming under field conditions are not found in the literature.
We conducted a field climate manipulative experiment and investigated developmental and demographic responses to diel asymmetric warming in three grasshopper species (an early-season species Dasyhippus barbipes, a mid-season species Oedaleus asiaticus, and a late-season species Chorthippus fallax). It was found that warming generally advanced the development of eggs and nymphs, but had no apparent impacts on the hatching rate of eggs, the emergence rate of nymphs and the survival and fecundity of adults in all the three species. Nighttime warming was more effective in advancing egg development than the daytime warming. The emergence time of adults was differentially advanced by warming in the three species; it was advanced by 5.64 days in C. fallax, 3.55 days in O. asiaticus, and 1.96 days in D. barbipes. This phenological advancement was associated with increases in the effective GDDs accumulation.
Results in this study indicate that the responses of the three grasshopper species to warming are influenced by several factors, including species traits, developmental stage, and the thermal sensitivity of the species. Moreover, species with diapausing eggs are less responsive to changes in temperature regimes, suggesting that development of diapausing eggs is a protective mechanism in early-season grasshopper for avoiding the risk of pre-winter hatching. Our results highlight the need to consider the complex relationships between climate change and specificity responses of invertebrates.
Although mutualism between ants and honeydew-producing hemipterans has been extensively recognized in ecosystem biology, however few attempts to test the hypothesis that mutualism between two alien species leads to the facilitation of the invasion process. To address this problem, we focus on the conditional mutualism between S. invicta and P. solenopsis by field investigations and indoor experiments. In the laboratory, ant colony growth increased significantly when ants had access to P. solenopsis and animal-based food. Honeydew produced by P. solenopsis also improved the survival of ant workers. In the field, colony density of P. solenopsis was significantly greater on plots with ants than on plots without ants. The number of mealybug mummies on plants without fire ants was almost three times that of plants with fire ants, indicating a strong effect of fire ants on mealybug survival. In addition, the presence of S. invicta successfully contributed to the spread of P. solenopsis. The quantity of honeydew consumption by S. invicta was significantly greater than that of a presumptive native ant, Tapinoma melanocephalum. When compared with the case without ant tending, mealybugs tended by ants matured earlier and their lifespan and reproduction increased. T. melanocephalum workers arrived at honeydew more quickly than S. invicta workers, while the number of foraging S. invicta workers on plants steadily increased, eventually exceeding that number of T. melanocephalum foragers. Overall, these results suggest that the conditional mutualism between S. invicta and P. solenopsis facilitates population growth and fitness of both species. S. invicta tends to acquire much more honeydew and drive away native ants, promoting their predominance. These results suggest that the higher foraging tempo of S. invicta may provide more effective protection of P. solenopsis than native ants. Thus mutualism between these two alien species may facilitate the invasion success of both species.
Scarcity of essential nutrients has led plants to evolve alternative nutritional strategies, such as myrmecotrophy (ant-waste-derived nutrition) and carnivory (invertebrate predation). The carnivorous plant Nepenthes bicalcarata grows in the Bornean peatswamp forests and is believed to have a mutualistic relationship with its symbiotic ant Camponotus schmitzi. However, the benefits provided by the ant have not been quantified. We tested the hypothesis of a nutritional mutualism, using foliar isotopic and reflectance analyses and by comparing fitness-related traits between ant-inhabited and uninhabited plants. Plants inhabited by C. schmitzi produced more leaves of greater area and nitrogen content than unoccupied plants. The ants were estimated to provide a 200% increase in foliar nitrogen to adult plants. Inhabited plants also produced more and larger pitchers containing higher prey biomass. C. schmitzi-occupied pitchers differed qualitatively in containing C. schmitzi wastes and captured large ants and flying insects. Pitcher abortion rates were lower in inhabited plants partly because of herbivore deterrence as herbivory-aborted buds decreased with ant occupation rate. Lower abortion was also attributed to ant nutritional service. The ants had higher δ15N values than any tested prey, and foliar δ15N increased with ant occupation rate, confirming their predatory behaviour and demonstrating their direct contribution to the plant-recycled N. We estimated that N. bicalcarata derives on average 42% of its foliar N from C. schmitzi wastes, (76% in highly-occupied plants). According to the Structure Independent Pigment Index, plants without C. schmitzi were nutrient stressed compared to both occupied plants, and pitcher-lacking plants. This attests to the physiological cost of pitcher production and poor nutrient assimilation in the absence of the symbiont. Hence C. schmitzi contributes crucially to the nutrition of N. bicalcarata, via protection of assimilatory organs, enhancement of prey capture, and myrmecotrophy. This combination of carnivory and myrmecotrophy represents an outstanding strategy of nutrient sequestration.
In the UK, attempts since the 1970s to control the incidence of bovine tuberculosis (bTB) in cattle by culling a wildlife host, the European badger (Meles meles), have produced equivocal results. Culling-induced social perturbation of badger populations may lead to unexpected outcomes. We test predictions from the ‘perturbation hypothesis’, determining the impact of culling operations on badger populations, movement of surviving individuals and the influence on the epidemiology of bTB in badgers using data dervied from two study areas within the UK Government's Randomised Badger Culling Trial (RBCT). Culling operations did not remove all individuals from setts, with between 34–43% of badgers removed from targeted social groups. After culling, bTB prevalence increased in badger social groups neighbouring removals, particularly amongst cubs. Seventy individual adult badgers were fitted with radio-collars, yielding 8,311 locational fixes from both sites between November 2001 and December 2003. Home range areas of animals surviving within removed groups increased by 43.5% in response to culling. Overlap between summer ranges of individuals from Neighbouring social groups in the treatment population increased by 73.3% in response to culling. The movement rate of individuals between social groups was low, but increased after culling, in Removed and Neighbouring social groups. Increased bTB prevalence in Neighbouring groups was associated with badger movements both into and out of these groups, although none of the moving individuals themselves tested positive for bTB. Significant increases in both the frequency of individual badger movements between groups and the emergence of bTB were observed in response to culling. However, no direct evidence was found to link the two phenomena. We hypothesise that the social disruption caused by culling may not only increase direct contact and thus disease transmission between surviving badgers, but may also increase social stress within the surviving population, causing immunosuppression and enhancing the expression of disease.