Fight or flight? A geographic mosaic in host reaction and potency of a chemical weapon in the social parasite <Emphasis Type="Italic">Harpagoxenus sublaevis</Emphasis>

Ant social parasites use chemical warfare to facilitate host colony takeover, which is a critical but recurring step in their life cycle. Many slave-making ants use the secretion of the Dufour gland to manipulate host behaviour during parasitic nest foundation and slave raids. Harpagoxenus sublaevis applies this chemical weapon onto defending Leptothorax host workers, which elicits deadly fights amongst them. Host species are expected to evolve counter-adaptations against this behavioural manipulation and in this study we investigated the geographic structure of this co-evolving trait. We compared the effectiveness of the parasitic gland secretion from different H. sublaevis populations in host colonies from various sites and analysed the occurrence of local adaptation. The two host species L. muscorum and L. acervorum generally showed different responses to the parasites’ chemical weapon: L. acervorum attacked nestmates treated with Dufour gland secretion, while L. muscorum workers fled. Flight, instead of intraspecific fights, is an adaptive host reaction as it results in fewer host fatalities during raids. Beside interspecific host differences, we found a geographic mosaic of host resistance: parasites from a German population strongly manipulated the behaviour of both sympatric Leptothorax populations. Russian or Italian hosts instead did not react with intracolonial aggression, but fled when confronted with the gland secretion of their sympatric parasite. Not only variation in host resistance explains differences in the effectiveness of the parasitic gland secretion but also interpopulational differences in its chemical composition, which were revealed by gas chromatography and mass spectrometry.     

Effects of parasitic mites and protozoa on the flower constancy and foraging rate of bumble bees

Parasites can affect host behavior in subtle but ecologically important ways. In the laboratory, we conducted experiments to determine whether parasitic infection by the intestinal protozoan Crithidia bombi or the tracheal mite Locustacarus buchneri alters the foraging behavior of the bumble bee Bombus impatiens. Using an array of equally rewarding yellow and blue artificial flowers, we measured the foraging rate (flowers visited per minute, flower handling time, and flight time between flowers) and flower constancy (tendency to sequentially visit flowers of the same type) of bees with varying intensities of infection. Bumble bee workers infected with tracheal mites foraged as rapidly as uninfected workers, but were considerably more constant to a single flower type (yellow or blue). In contrast, workers infected with intestinal protozoa showed similar levels of flower constancy, but visited 12% fewer flowers per minute on average than uninfected bees. By altering the foraging behavior of bees, such parasites may influence interactions between plants and pollinators, as well as the reproductive output of bumble bee colonies. Our study is the first to investigate the effects of parasitic protozoa and tracheal mites on the foraging behavior of bumble bees, and provides the first report of Crithidia bombi in commercial bumble bees in North America.     

Division of labor and slave raid initiation in slave-making ants

In social insect societies, division of labor, i.e., workers of a colony specializing in different tasks, is thought to improve colony performance. Workers of social parasitic slave-making ants focus on a single task, searching for and raiding host colonies to replenish their slave workforce. However, in the North American slavemaker Protomognathus americanus, some workers do not partake in raids but remain inside the colony. We analyzed raid participation, fertility, and cuticular hydrocarbon profiles of slavemaker workers and slaves to understand these behavioral differences and the regulation of division of labor in slavemaker colonies. Raid observations showed that some workers were repeatedly involved in raiding activities (exterior workers), whereas others stayed inside the nest (interior workers). Exterior workers were always infertile, while half of the interior workers were fertile. Analysis of cuticular hydrocarbons demonstrated differences between the groups. We also detected chemical differences between interior and exterior slaves, indicating an influence of the individuals’ tasks on their cuticular profiles. Task- and fertility-related profiles may allow selective nestmate recruiting. Division of labor should also adapt to varying conditions. Since slave raids are dangerous, they should only be initiated when the colony needs additional slaves. Exclusively fed by their slaves, slavemaker workers could determine this need via their nutritional status. In an experiment with various feeding regimes, colonies subjected to a lower food provisioning rate showed increased proportions of slavemaker workers searching for host colonies. Division of labor in slave-making ants, therefore, might be flexible and can change depending on the colonies’ needs.     

Parasitism in a social wasp: effect of gregarines on foraging behavior, colony productivity, and adult mortality

Behavior in eusocial insects likely reflects a long history of selection imposed by parasites and pathogens because the conditions of group living often favor the transmission of infection among nestmates. Yet, relatively few studies have quantified the effects of parasites on both the level of individual colony members and of colony success, making it difficult to assess the relative importance of different parasites to the behavioral ecology of their social insect hosts. Colonies of Polybia occidentalis, a Neotropical social wasp, are commonly infected by gregarines (Phylum Apicomplexa; Order Eugregarinida) during the wet season in Guanacaste, Costa Rica. To determine the effect of gregarine infection on individual workers in P. occidentalis, we measured foraging rates of marked wasps from colonies comprising both infected and uninfected individuals. To assess the effect of gregarines on colony success, we measured productivity and adult mortality rates in colonies with different levels of infection prevalence (proportion of adults infected). Foraging rates in marked individuals were negatively correlated with the intensity of gregarine infection. Infected colonies with high gregarine prevalence constructed nests with fewer brood cells per capita, produced less brood biomass per capita, and, surprisingly, experienced lower adult mortality rates than did uninfected or lightly infected colonies. These data strongly suggest that gregarine infection lowers foraging rates, thus reducing risk to foragers and, consequently, reducing adult mortality rates, while at the same time lowering per-capita input of materials and colony productivity. In infected colonies, queen populations were infected with a lower prevalence than were workers. Intra-colony infection prevalence decreased dramatically in the P. occidentalis population during the wet season.An erratum to this article can be found at     

Population and seasonal variation in response to prey calls by an eavesdropping bat

The fringe-lipped bat, Trachops cirrhosus, is an eavesdropping predator that hunts frogs and katydids by approaching these preys' sexual advertisement calls. In captivity, bats can rapidly learn to associate novel acoustic stimuli with food rewards. It is unknown how this learning ability is related to foraging behavior in the wild where prey and the calls that identify them vary over space and time. In two bat populations that differ in available prey species (Soberanía, Panama, and La Selva, Costa Rica), we presented wild-caught bats with frog calls, katydid calls, and control stimuli. Bats in Soberanía were significantly more responsive to complex calls and choruses of the túngara frog, Physalaemus pustulosus, than were bats in La Selva. La Selva bats were significantly more responsive to katydid calls (Steirodon sp.) than Soberanía bats. We also examined seasonal variation in bat response to prey cues. Bats were captured in Soberanía in dry and wet seasons and presented with the calls of a dry season breeding frog (Smilisca sila), a wet season breeding frog (P. pustulosus), and four katydid species. Bats captured in the dry season were significantly more responsive to the calls of S. sila than bats captured in the wet season, but there were no seasonal differences in response to the calls of P. pustulosus or the katydid calls. We demonstrate plasticity in the foraging behavior of this eavesdropping predator but also show that response to prey cues is not predicted solely by prey availability.     

Seagrass and algal beds as nursery habitats for tiger prawns (Penaeus semisulcatus and P. esculentus) in a tropical Australian estuary

We evaluated the importance of seagrass and algae to two species of tiger prawns (Penaeus semisulcatus and P. esculentus) by detailed sampling at four sites (two seagrass, two algae) in the Embley River estuary, and through sampling 26 sites in 7 adjacent estuaries at one time of year. Samples of tiger prawns were collected in the Embley River estuary with a small beam trawl at night every 2 wk from September to May for 2 yr (1990 to 1992). The two seagrass sites, which were 11 and 13 km from the river mouth, showed less seasonal variation in salinity than the two algal sites, which were 15 and 20 km from the river mouth. The algal beds at the two upstream sites almost disappeared during the wet season, but the biomass of seagrass did not change significantly between the wet and dry seasons. The grooved tiger prawn (P. semisulcatus), the main species at all sites, comprised 88% of the total tiger prawn catch over the two years. They were found at all sites during the pre-wet season, but after the onset of the wet season, they disappeared along with the algae, from the upstream sites. The brown tiger prawn (P. esculentus) was found almost exclusively (97% of the total catch) on the seagrass sites downstream. In the study of several estuaries, juvenile P. semisulcatus were caught at all 26 sites, and P. esculentus were caught in much smaller numbers, at 16 sites. Approximately equal numbers of P. semisulcatus were caught in seagrass and algal beds in the pre-wet season. Very few individuals >10 mm carapace length of either species, were caught. The results from this study highlight the importance of algal beds during the pre-wet season as nursery areas for one species of tiger prawn (P. semisulcatus).     

Vibrational communication between hitchhikers and foragers in leaf-cutting ants (Atta cephalotes)

Abstract In a foraging column of the leaf-cutting ant Atta cephalotes, minim workers (the smallest worker subcaste) hitchhike on leaf fragments carried by larger workers. It has been demonstrated that they defend leaf carriers against parasitic phorid flies. The present study examines the cues used by the potential hitchhikers to locate leaf carriers. As recently reported, foraging workers stridulate while cutting a leaf fragment, and the stridulatory vibrations serve as closerange recruitment signals. We tested the hypothesis that these plant-borne stridulatory vibrations are used by the potential hitchhikers to locate workers engaged in cutting. Three different lines of evidence support this view. Firstly, the repetition rate of the stridulations produced by foraging workers increases significantly as foragers maneuver the leaf fragment into the carrying position and walk loaded to the nest. This is the moment when hitchhikers usually climb on the leaf. Although the leaf-borne stridulatory vibrations are considerably attenuated when transmitted through the workers' legs, they can nevertheless be detected at short distances by minims. This subcaste is several times more sensitive to substrate-borne vibrations than larger workers. Secondly, when a stridulating and a silent leaf were simultaneously presented at the foraging site, minim workers spent significantly more time on the stridulating than on the silent leaf. Thirdly, hitchhiking was more frequent in leaf carriers which cut fragments out of the stridulating leaf than in those cutting the silent leaf.Abstract In a foraging column of the leaf-cutting ant Atta cephalotes, minim workers (the smallest worker subcaste) hitchhike on leaf fragments carried by larger workers. It has been demonstrated that they defend leaf carriers against parasitic phorid flies. The present study examines the cues used by the potential hitchhikers to locate leaf carriers. As recently reported, foraging workers stridulate while cutting a leaf fragment, and the stridulatory vibrations serve as closerange recruitment signals. We tested the hypothesis that these plant-borne stridulatory vibrations are used by the potential hitchhikers to locate workers engaged in cutting. Three different lines of evidence support this view. Firstly, the repetition rate of the stridulations produced by foraging workers increases significantly as foragers maneuver the leaf fragment into the carrying position and walk loaded to the nest. This is the moment when hitchhikers usually climb on the leaf. Although the leaf-borne stridulatory vibrations are considerably attenuated when transmitted through the workers' legs, they can nevertheless be detected at short distances by minims. This subcaste is several times more sensitive to substrate-borne vibrations than larger workers. Secondly, when a stridulating and a silent leaf were simultaneously presented at the foraging site, minim workers spent significantly more time on the stridulating than on the silent leaf. Thirdly, hitchhiking was more frequent in leaf carriers which cut fragments out of the stridulating leaf than in those cutting the silent leaf.Communicated by P. Pamilo     

Defense against parasites by hitchhikers in leaf-cutting ants: a quantitative assessment

Summary A curious behavior in leaf-cutting ants in the genus Atta is the hitchhiking of small minim workers on leaf fragments carried by larger workers. Two functions of these hitchhikers have been proposed: (1) defense of leaf carriers against parasitic flies in the family Phoridae (ant protection hypothesis; Eibl-Eibesfeldt and Eibl-Eibesfeldt 1967) and (2) reduction of transport costs of small minims that collect plant fluids (energy conservation hypothesis; Stradling 1978).We studied hitchhiking behavior in colonies of Atta colombica on Barro Colorado Island, Panama, and found strong evidence in favor of the ant protection hypothesis. Females of Apocephalus attophilus (Diptera: Phoridae) attack leaf carriers of A. colombica and deposit eggs in the head capsules of these ants. Our observations indicate that parasites require leaf fragments to stand on during oviposition, and, as a result, only leaf carriers are susceptible to parasitic attack. The presence of hitchhikers reduces significantly both the time parasites spend on leaf fragments and the probability that they will land in the first place. Results of experimental introductions of parasites and a year of biweekly censuses at ten colonies indicate that leaf-cutting ants adjust the level of hitchhiking to accommodate both daily and seasonal changes in the abundance of parasites.We found little evidence in support of the energy conservation hypothesis. If it is assumed that all minim workers hitchhike back to the nest, our calculations indicate that total transport costs along a foraging trail are reduced by 10% or less. However, our observations indicate that only 50% of returning minim workers hitchhike, and therefore energy savings are actually considerably less than 10%. Leaf-cutting ants in the genera Atta and Acromyrmex are attacked by over 20 species of parasitic phorids. In the discussion we review what is known about these associations and suggest that these parasites have influenced the ecology and evolution of polyethism in leaf-cutting ants. Offprint requests to: D.H. Feener (at his present address)     

Co-evolution-driven cuticular hydrocarbon variation between the slave-making ant Rossomyrmex minuchae and its host Proformica longiseta (Hymenoptera: Formicidae)

Summary. Proformica longiseta exists as two populations in the Sierra Nevada Mountains in Spain, only one of which is parasitized by the slave-maker ant Rossomyrmex minuchae. To investigate the possible effect of co-evolutionary pressures on cuticular hydrocarbon (CHC) profiles (the presumed nestmate recognition cues), we performed a comparative analysis of the CHC of R. minuchae and P. longiseta colonies from both the allopatric and sympatric populations; the latter includes samples of enslaved as well as free-living workers. Discriminant analyses based on these chemical profiles showed two clear profile groups: the first comprised R. minuchae and both enslaved and free-living P. longiseta from the sympatric population; and the second the allopatric P. longiseta workers. As expected, the profiles of the two sympatric P. longiseta groups (enslaved and free-living) were distinct; but, interestingly, those of the enslaved P. longiseta and its parasite R. minuchae were also distinguishable. This indicates that despite their cohabitation each species maintains its own chemical identity. Profile similarity between the sympatric free-living P. longiseta and its parasite may explain the lower than expected aggression observed during raids. We further speculate that in view of the differences between the sympatric and allopatric population of P. longiseta, co-evolutionary pressures have driven changes in the profile of the former to better match that of its parasite R. minuchae. Such an adjustment may have enabled nests of the sympatric P. longiseta to endure multiple raids by the parasite (due to the reduced aggression) and thus to continue to reproduce despite the damage inflicted by the raids.     

Do these eggs smell funny to you?: an experimental study of egg discrimination by hosts of the social parasite Polyergus breviceps (Hymenoptera: Formicidae)

Social parasites exploit the behaviours of other social species. Infiltration of host systems involves a variety of mechanisms depending on the conditions within the host society and the needs of the social parasite. For many species of socially parasitic ants, colony establishment entails the usurpation of colonies of other species. This frequently involves the eviction or death of the host colony queen and the subsequent adoption of the invading queen. The social parasite queen achieves host worker acceptance by either manipulating the nest-mate recognition processes of the host or undergoing chemical modification. Little is known, however, about how host workers respond to social parasite eggs or whether host species defend against brood parasitism during parasite invasions. Host species are believed to adopt social parasite offspring because the recent common ancestry between many social parasites and their hosts may grant the sharing of certain characteristics such as chemical cues. Use of multiple host species, however, suggests other processes are needed for the social bond between host and parasite young to form. This study reports the findings of adoption bioassays in which eggs from a slave-maker ant, Polyergus breviceps, were offered to workers of two of its host species from unparasitised or newly parasitised nests to determine whether P. breviceps eggs generally elicit rearing behaviours from multiple host species. Comparisons of parasite egg survival until adulthood with conspecific egg survival reveal that workers of both host species, free-living or newly enslaved, do not typically accept slave-maker eggs. Both host species thus have sufficient discriminatory power to reject social parasite eggs although our hydrocarbon analysis indicates parasite eggs may be adapted to their local host species. Combined these results suggest that host rearing of P. breviceps eggs may reflect an evolutionary equilibrium that is maintained by probability and cost of recognition errors.Communicated by L. Sundström     

Phytoextraction potential of Sporobulus pyramidalis found in an abandoned battery industry environment in Nigeria

The levels of heavy metals (Cd, Cr, Ni, Pb, Sb and V) in Sporobulus pyramidalis plant species from an abandoned battery industry environment were determined during the wet and dry seasons as follows: The ranges in metal concentrations during the wet and dry seasons were 0.002, 0.420, 0.036 and 36.10?µg?g^?1, respectively. The coefficients of variation for the metals during wet and dry seasons ranged between none to 151.724% and 13.838–214.935%, respectively. The results showed higher levels of the metals in S. pyramidalis during the dry than wet season. Results obtained from both wet and dry season when compared with background values, Federal Environmental Protection Agency and other international standards revealed that the plant species accumulated high levels of these heavy metals which was evident in concentrations exceeding maximum tolerable limits. The health implications of consuming this plant and any other plant or crop within this environment are discussed.     

A South East Asian ponerine ant of the genus Leptogenys (Hym., Form.) with army ant life habits

Summary The emigration and raiding behavior of the SE Asian ponerine ant Leptogenys sp. 1, which resembles L. mutabilis, were observed in the field (Ulu Gombak, Malaysia). The ants formed monogynous colonies that consisted of up to 52 100 workers. The bivouac sites of this species were found in leaf litter, rotten logs, ground cavities, etc., and were rarely modified by the ants. The colonies stayed in these temporary nests for several hours to 10 days; afterwards, they moved to a new nest site. The emigration distances ranged from 5–58 m. Since nest changing takes place at irregular intervals, and pupae and larvae are always present in the nest relocations of Leptogenys sp. 1, the emigration behavior is not linked to a synchronized brood development. Leptogenys sp. 1 is a nocturnal forager; in our study, up to 42 600 workers participated in each raid. The ants move forward on a broad front; behind the swarm a fan-shaped network of foraging columns converges to form a main trunk trail. A new system of foraging trails is developed in each raid. The workers search for their prey collectively; they attack and retrieve the booty together. The diet of Leptogenys sp. 1 consists mainly of arthropods. Army ant behavior is characterized by (1) formation of large monogynous colonies, (2) frequent emigrations, and (3) mass raids in which all foraging activities are carried out collectively. Since Leptogenys sp. 1 performs these typical army ant behavior patterns, this species represents the army ant ecotype. However, this species differs considerably from army ant species that have synchronized broods and huge colonies with dichthadiiform queens.Dedicated to Professor Dr. M. Lindauer on the occasion of his 70th birthday     

Social parasitism by honeybee workers (Apis mellifera capensis Esch.): evidence for pheromonal resistance to host queen’s signals

Social parasites exploit their host’s communication system to usurp resources and reproduce. In the honeybee, Apis mellifera, worker reproduction is regulated by pheromones produced by the queen and the brood. Workers usually reproduce when the queen is removed and young brood is absent. However, Cape honeybee workers, Apis mellifera capensis, are facultative intraspecific social parasites and can take over reproduction from the host queen. Investigating the manner in which parasitic workers compete with host queens pheromonally can help us to understand how such parasitism can evolve and how reproductive division of labour is regulated. In A. m. capensis, worker reproduction is associated with the production of queen-like pheromones. Using pheromonal contest experiments, we show that Apis mellifera scutellata queens do not prevent the production of queen-like mandibular gland compounds by the parasites. Given the importance of these pheromones in acquiring reproductive status, our data suggest that the single invasive lineage of parasitic workers occurring in the range of A. m. scutellata was selected for its superior ability to produce these signals despite the presence of a queen. Such resistance was indeed less frequent amongst other potentially parasitic lineages. Resistance to reproductive regulation by host queens is probably the key factor that facilitates the evolution of social parasitism by A. m. capensis workers. It constitutes a mechanism that allows workers to evade reproductive division of labour and to follow an alternative reproductive option by acquiring direct fitness in foreign colonies instead of inclusive fitness in their natal nests.     

Parasitic flies (Diptera: Phoridae) influence foraging rhythms and caste division of labor in the leaf-cutter ant,Atta cephalotes (Hymenoptera: Formicidae)

Summary Three lines of evidence, including interspecific comparisons, temporal division of foraging between size castes, and experimental manipulations, suggest that the diurnal parasitoid Neodohrniphora curvinervis (Diptera: Phoridae) influences both the caste sizes and numbers of leaf-cutter ants (Atta cephalotes) that leave their underground nests to collect leaves. At Parque Nacional Corcovado in Costa Rica, A. cephalotes was attacked by Neodohrniphora during the daytime, and foraged less during the day than at night; a closely related ant at the same site, A. colombica, had no phorid parasites and foraged exclusively during the day. Most daytime foragers of A. cephalotes were smaller than the lower size threshold for attack by Neodohrniphora, while nocturnal foragers, active when parasitoids were absent, were both larger than this threshold and within the energetically optimal size range for foraging. When I supplied artificial lighting to allow phorids to hunt at A. cephalotes colonies past dusk, ants foraged less than when light was provided but flies were removed. The influence of Neodohrniphora on the foraging activity of A. cephalotes may explain why investigations focusing on abiotic factors have largely failed to discover what drives this ant's daily foraging cycles, and suggests that forager sizes are influenced not only by energetic efficiency, but also by the threat of parasitism.     

Chemical camouflage of the slave-making ant Polyergus samurai queen in the process of the host colony usurpation (Hymenoptera: Formicidae)

Founding queens of the obligatory social parasite ant Polyergus samurai usurp the host ant Formica japonica colony. The aggressive behaviors of F. japonica workers on the parasite queen disappear after the parasite queen kills the resident queen. To determine whether the parasite queen chemically mimics the host ants, we examined the aggressive behavior of F. japonica workers toward glass dummies applied with various extracts of the parasite queen and host workers. The crude extracts and hydrocarbon fraction reproduced the host workers’ behavior to the live ants. The extracts of the post-adoption parasite queen, as well as the nestmate extracts of F. japonica, did not elicit the aggressive behavior, but the extract of the pre-adoption parasite queen triggered attacks by the host workers. The nestmate recognition of host workers did not change, regardless of contact with the parasite. The gas chromatography and gas chromatography–mass spectrometry analyses indicated that the cuticular hydrocarbon (CHC) profile of the parasite queen drastically changed during the process of usurpation. Discriminant analysis showed the successfully usurped P. samurai queen had colony-specific CHC profiles. CHC profiles of the P. samurai queen who killed the host queen were more similar to those of the host queen than the workers, while the P. samurai queen who usurped the queenless colony had a profile similar to those of host workers. These results suggest that the P. samurai queen usually acquires the CHCs from the host queen during the fight, but from host wokers in queenless host colonies.     

Activity patterns of giant petrels, Macronectes spp., using different foraging strategies

We studied foraging activity of giant petrels during the incubation period, by simultaneously deploying activity recorders and satellite transmitters on northern (Macronectes halli) and southern giant petrels (Macronectes giganteus) at Bird Island (South Georgia, Antarctica) between 29 October and 26 December 1998. Satellite tracking showed two types of trips: (1) coastal trips, all undertaken by male northern giant petrels, to the nearby South Georgia mainland, presumably foraging on seal and penguin carcasses on beaches, and (2) pelagic trips, foraging at sea for marine prey or potentially scavenging on distant archipelagos (e.g. South Sandwich, Falkland or South Orkney Islands). Activity recorder data were consistent with the types of trip defined by the satellite tracking data, with median wet activity (time spent at the sea surface) during pelagic trips being 41%, but only 14% on coastal trips. On pelagic trips, there was a significant negative correlation between the duration of wet periods and the speed of travel between satellite uplinks. Mean travelling speed between uplinks was greater during day than night for both types of trips, suggesting that giant petrels prefer to travel during daylight and are less active at night. The scarcity of wet periods during the night in giant petrels foraging to the South Georgia coast (median=3%, range=1-9%) indicates that such birds spent almost all night on land. Likewise, the scarcity of wet periods at night for three birds foraging 700-1,000 km south of Bird Island, where there is no land but abundant icebergs, suggests these birds were resting on the icebergs at night. In addition to the adaptations to scavenging on carrion, pelagic trips by giant petrels contain elements similar to those of albatrosses, indicating a complexity to giant petrel lifestyle hitherto unrecognised.     

Seasonal variations in trace metal levels,speciation and physicochemical determinants of metal availability in dumpsite soils within Akwa Ibom State,Nigeria

Physicochemical properties, total metals and metal species were investigated in surface soil samples from six urban dumpsites within Akwa Ibom State, Nigeria. Results obtained indicated that Pb, Cd and Cr levels were higher in the dry season than in wet season, whereas Ni levels were higher during the wet season. Soil pH, organic matter and cation-exchange capacity (CEC) showed higher values in wet season than in dry season, whereas sand and clay levels were higher in the dry season. Speciation analysis showed the principal fraction of the metals to be Pb (reducible), Cd (acid extractable) and Ni (residual), whereas Cr existed in the residual fraction during the dry season and in the acid-extractable fraction during the wet season. Soil pH was positively correlated at p<0.1 with all metals in both seasons, sand was negatively correlated with the metals in both seasons, and CEC was negatively correlated with all metals in both seasons, except Pb in the wet season. Organic matter showed variable relationships with metals in both seasons. Seasonal variations affected metal availability and physicochemical properties of the soil. Total metal concentrations followed the order Pb>Ni>Cd>Cr, whereas their bioavailability followed the trend Cd>Pb>Cr>Ni.     

Thermal ecology,size effects,and the origins of communal behaviour in Cerceris wasps

Summary Female Cerceris nest in large aggregations in sandy soil, stocking cells with weevils to feed the offspring. Their foraging patterns depend upon weather conditions, and the timing and duration of foraging is predictably variable according to their size. Hence large foragers collect more prey, and can stock more cells, and through a normal season they should stock twice as many cells (thus laying twice as many eggs) as small females. Since they also stock more prey per cell, and provisioning determines offspring size, large females thus produce more and larger children simply through the effects of the weather. Despite their apparent disadvantages, small female wasps never-theless persist, and various factors contribute to this; they do better in hot weather, when large females do not forage due to over-heating risks, they tend to bias their sex ratio more to male production, and they often co-occupy a nest with a larger female. Nest co-occupancy increases in very hot conditions, when a large female benefits by the foraging of a small one while it is nest-bound, a small female benefits by getting a chance to lay in better-stocked cells, and both may benefit from decreased parasitism by satellite flies. Thus two factors which may contribute to joint nesting and a trend towards eusociality along the parasocial route are suggested. Firstly such a trend is more likely in hot dry conditions; and secondly it is more likely in species with a broad range of female sizes. Both these effects can be seen in a survey of wasps of the genus Cerceris so far reported in the literature, and are likely to be more widely applicable in studies where weather effects interacting with foraging patterns have hitherto been ignored.     

Contrasting home-range size and spatial partitioning in cryptic and sympatric pipistrelle bats

The extent of spatial partitioning in insectivorous bats, whose prey is patchily distributed and transient in nature, remains a contentious issue. The recent separation of a common Palaearctic bat, the pipistrelle, into Pipistrellus pipistrellus and Pipistrellus pygmaeus, which are morphologically similar and sympatric, provides an opportunity to examine this question. The present study used radio telemetry to address the spatial distribution and foraging characteristics of P. pipistrellus and P. pygmaeus in northeast Scotland, to test the hypothesis that coexistence between these species is facilitated through spatial segregation. We reveal large and significant differences in the spatial distribution and foraging characteristics of these two cryptic species. Individual P. pipistrellus home ranges were on average three times as large as that of P. pygmaeus, and they foraged for approximately an hour longer each night. Inter-specific spatial overlap was minimal (<5%) and core foraging areas of either species were essentially mutually exclusive despite the proximity of the two roosts. Inter-specific differences in range size were associated with the spatial dispersion of productive foraging sites within individual foraging ranges. P. pipistrellus foraging sites were highly dispersed, necessitating larger ranges. It is predicted that the spatial segregation revealed by the present study is a result of selection favouring the avoidance of competition in these species through differential habitat use.     

Lek promiscuity in a semelparous mammal,Antechinus stuartii (Marsupialia: Dasyuridae)?

Summary All male Antechinus stuartii die following a brief rut towards the end of their first year of life. Outside the rutting period, both males and females forage in clearly-defined invidual home ranges, but neither sex is territorial. As temperatures drop during the winter, both sexes may leave their foraging range each day to spend the latter part of the night and most of the daylight hours in communal nests which may be used by fifteen animals simultaneously, and over forty individuals during one winter. These temporary movements away from the foraging range cause an increase in the daily home range revealed by radiotelemetry. When the mating season commences, males abandon their foraging range and aggregate in a few of the communal nest trees, where they spend most of the night. There is some movement between aggregations, but the movement appears to be rapid and direct. Females continue to use their foraging area, and nest solitarily. However, they make excursions to the male aggregations, and spend time in the nests with the males. By the third week of the mating season, immediately prior to male mortality, males move frequently from one aggregation to another, but are still confined to the aggregations and direct movements between those nests. By this time some females have ceased visiting the aggregations and nest solitarily within their foraging range. These data suggest that the mating system of A. stuartii is lek promiscuity, and support the hypothesis that the abrupt mortality of males is a cost of endocrine changes which facilitate gluconeogenic mobilisation of body protein, allowing the males to sustain their vigil at the leks.