Queen recruitment and split sex ratios in polygynous colonies of the ant Formica exsecta

Sex ratios in social insects have become a general model for tests of inclusive fitness theory, sex ratio theory and parent–offspring conflict. In populations of Formica exsecta with multiple queens per colony , sex ratios vary greatly among colonies and the dry-weight sex ratio is extremely male-biased, with 89% of the colonies producing males but no gynes (reproductive females). Here we test the queen-replenishment hypothesis, which was proposed to explain sex ratio specialization in this and other highly polygynous ants (i.e. those with many queens per nest). This hypothesis proposes that, in such ants, colonies produce gynes to recruit them back into the colony when the number of resident queens falls below a given threshold limiting colony productivity or survival. We tested predictions of the queen-replenishment hypothesis by following F. exsecta colonies across two breeding seasons and relating the change in effective queen number with changes in sex ratio, colony size and brood production. As predicted by the queen-replenishment hypothesis, we found that colonies that specialized in producing females increased their effective queen number and were significantly more likely to specialize in male production the following year. The switch to male production also coincided with a drop in productivity per queen as predicted. However, adoption of new queens did not result in a significant increase in total colony productivity the following year. We suggest that this is because queen production comes at the expense of worker production and thus queen production leads to resource limitation the following year, buffering the effect of greater queen number on total productivity.     

Varying foraging strategies of Labridae in seagrass habitats: Herbivory in temperate seagrass meadows?

The diets of five species of Labridae in south-western Australia were examined to determine whether: (1) grazing of seagrass and epiphytic algae is a prominent feature of the food web within the deeper seagrass meadows of this temperate region; (2) levels of grazing differ among different seagrass systems; and diets differ among these closely-related species. Fish were collected seasonally from three seagrass habitats mainly comprising either Posidonia sinuosa, Posidonia coriacea or Amphibolis griffithii between the summer of 1996/97 and spring of 1997. Consumption of considerable amounts of algae and seagrass by Odax acroptilus and seagrass by Haletta semifasciata indicates that macrophyte grazing by fish is a component of the trophic dynamics of south-western Australian seagrass meadows. O. acroptilus and H. semifasciata were both omnivorous, feeding on a range of epifauna, infauna and flora, whereas Siphonognathus radiatus, Neoodax balteatus and Notolabrus parilus were carnivorous, feeding predominantly on motile epifauna, such as molluscs and crustaceans. The level of macrophyte grazing is likely to be underestimated in temperate offshore meadows of P. sinuosa and A. griffithii where omnivorous labrids, monacanthids and terapontids are abundant. Stable isotope data for O. acroptilus from the study region suggest that animal prey is more important to tissue maintenance than macrophyte material. Macrophytes may be grazed to acquire attached animal prey or for fulfilling energy requirements. Based on the distribution of prey, it appears that species in A. griffithii meadows forage within and below the seagrass canopy, whilst species in P. sinuosa meadows are likely to forage towards the basal area of this seagrass.     

Inferring resource distributions from Atlantic bluefin tuna movements: an analysis based on net displacement and length of track

We use observed movement tracks of Atlantic bluefin tuna in the Gulf of Maine and mathematical modeling of this movement to identify possible resource patches. We infer bounds on the overall sizes and distribution of such patches, even though they are difficult to quantify by direct observation in situ. To do so, we segment individual fish tracks into intervals of distinct motion types based on the ratio of net displacement to length of track (ΔD/ΔL) over a time window Δt. To find the best segmentation, we optimize the fit of a random-walk movement model to each motion type. We compare results from two distinct movement models: biased turning and biased speed, to check the model-dependence of our inferences, and find that uncertainty in choice of movement model dominates the uncertainties of our conclusions. We find that our data are best described using two motion types: “localized” (ΔD/ΔL small) and “long-ranged” (ΔD/ΔL large). The biased turning model leads to significantly better resolution of localized movement intervals than the biased speed model. We hypothesize that localized movement corresponds to exploitation of resource patches. Comparison with visual behavior observations made during tracking suggests that many inferred intervals of localized motion do indeed correspond to feeding activity. From our analysis, we estimate that, on average, bluefin tuna in the Gulf of Maine encounter a resource patch every 2 h, that those patches have an average radius of 0.7-1.2 km, and that, overall, there are at most 5-9 such patches per in the region studied.     

Evolution of cooperation in spatial public goods games with common resource dynamics

Investment in a common resource shared by all players is difficult to evolve despite higher returns because a non-investor (free-rider) always receives more than an investor (altruist). This situation is referred to as the Tragedy of the Commons and is often observed in various biological systems including environmental problems of human society. Punishment and reputation are effective mechanisms but require cooperator's ability to identify free-riders. Volunteering can work in anonymous public goods games but this requires voluntary participation, which is not always the case. Here, we show that the evolution of altruism is possible in anonymous and obligate public goods games if we consider the spatiotemporal dynamics of the common resource that incorporate spatial diffusion and internal dynamics of the commons. The investors' strategy to counter free-riders is to increase population density and to outnumber them with the common resource level kept as low as that of the free-riders.     

Food supply modifies the trade-off between past and future reproduction in a sexual parasite–host system (<Emphasis Type="Italic">Rana esculenta,Rana lessonae</Emphasis>)

Life history theory is concerned with the costs of survival, growth and reproduction under different ecological conditions and the allocation of resources to meet these costs. Typical approaches used to address these topics include manipulation of food resources, followed by measures of subsequent reproductive traits, and measures of the relationship between current and future reproductive investment. Rarely, however, do studies test for the interaction of past investment, present resource availability and future investment simultaneously. Here, we investigate this interaction in females of a sexual parasite–host system consisting of the hybridogenetic frog Rana esculenta (E) and one of its parental species Rana lessonae (L). We kept females from each of two groups (with or without previous reproduction) under two food treatments (low or high) and regularly recorded their growth as well as their body condition and hormone titres as measures of future reproductive condition. After keeping them in hibernation until the following spring, we exposed the females to males, recorded whether they spawned or not and related this response to their condition in the previous autumn. Past reproduction negatively affected growth during summer and condition during autumn which, in turn, reduced the following year’s reproductive output. These costs of previous reproduction were less pronounced under the high than under the low food treatment and lower in R. lessonae than in R. esculenta. Increasing food supply improved reproductive condition more in L than in E females. These species differences in reproductive costs and food requirements provide a mechanistic explanation for why E females skip annual reproduction almost twice as often as L females. Since R. esculenta is a sexual parasite that depends on R. lessonae for successful reproduction, these species-specific life history patterns not only affect individual fitness but also the spatial structure and temporal dynamics of mixed LE populations.     

Size-dependent use of territorial space by a rock-dwelling cichlid fish

Territoriality fundamentally influences animal mating systems and patterns of population structure. Although territory ownership is already known to contribute importantly to male reproductive success and the ecological coexistence of African rock-dwelling cichlids, the significance of variation in territory features has received little attention in these fishes. In Lake Malawi, males of Pseudotropheus tropheops "orange chest" defend territories on either of two substrate classes at Harbour Island: flat rock slabs lacking crevices and caves, or structurally complex boulder fields containing cave shelters. Focal watches of this species demonstrated that both territory size and occupancy on either substrate type depend on the size of male residents. Males larger than a threshold size exclusively held the largest and most structurally complex territories. After removal of conspecific residents, more vacant territorial areas on cave-containing substrate were reoccupied by "orange chest" males in full breeding coloration compared to vacant areas on flat substrate. These findings suggest competition among "orange chest" males for complex rocky substrate. Defense of caves was associated with enhanced male courtship rates: the number of caves within a male's territory was a better predictor of courtship activity than was male size or territory area. In addition to territories being crucial for male reproductive success and therefore likely playing a role in sexual selection, male-male competition for caves in rock-dwelling cichlids may be promoted by the ecological advantage of enemy-free space. Smaller "orange chest" males lacking caves tended to move into adjacent boulder fields in the presence of predators, particularly at night. In contrast, males defending caves were more likely to remain on their territories when nocturnal predators were present. The territorial behaviors of P. tropheops "orange chest" that we observed in situ provide an instructive natural framework for testing the roles of substrate and ecology in the mating systems of rock-dwelling cichlid fishes.     

Multiscale wolf predation risk for elk: does migration reduce risk?

While migration is hypothesized to reduce predation risk for ungulates, there have been few direct empirical tests of this hypothesis. Furthermore, few studies examined multiscale predation risk avoidance by migrant ungulates, yet recent research reveals that predator–prey interactions occur at multiple scales. We test the predation risk reduction hypothesis at two spatial scales in a partially migratory elk (Cervus elaphus) population by comparing exposure of migrant and resident elk to wolf (Canis lupus) predation risk. We used GPS and VHF telemetry data collected from 67 migrant and 44 resident elk over the summers of 2002–2004 in and adjacent to Banff National Park (BNP), Canada. We used wolf GPS and VHF telemetry data to estimate predation risk as a function of the relative probability of wolf occurrence weighted by a spatial density model that adjusted for varying pack sizes. We validated the predation risk model using independent data on wolf-killed elk, and showed that combining wolf presence and spatial density best predicted where an elk was likely to be killed. Predation risk on summer ranges of migrant elk was reduced by 70% compared to within resident elk summer ranges. Because wolves avoided areas near high human activity, however, fine-scale selection by resident elk for areas near high human activity reduced their predation risk exposure to only 15% higher than migrants, a difference significant in only one of three summers. Finally, during actual migration, elk were exposed to 1.7 times more predation risk than residents, even though migration was rapid. Our results support the hypothesis that large-scale migrations can reduce predation. However, we also show that where small-scale spatial variation in predation risk exists, nonmigratory elk may equally reduce predation risk as effectively as migrants under some circumstances.     

Biomass responses to elevated CO<Subscript>2</Subscript>, soil heterogeneity and diversity: an experimental assessment with grassland assemblages

While it is well-established that the spatial distribution of soil nutrients (soil heterogeneity) influences the competitive ability and survival of individual plants, as well as the productivity of plant communities, there is a paucity of data on how soil heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. To evaluate the effects of elevated CO₂, soil heterogeneity and diversity (species richness and composition) on productivity, patterns of biomass allocation and root foraging precision, we conducted an experiment with grassland assemblages formed by monocultures, two- and three-species mixtures of Lolium perenne, Plantago lanceolata and Holcus lanatus. The experiment lasted for 90 days, and was conducted on microcosms built out of PVC pipe (length 38 cm, internal diameter 10 cm). When nutrients were heterogeneously supplied (in discrete patches), assemblages exhibited precise root foraging patterns, and had higher total, above- and belowground biomass. Greater aboveground biomass was observed under elevated CO₂. Species composition affected the below:aboveground biomass ratio and interacted with nutrient heterogeneity to determine belowground and total biomass. Species richness had no significant effects, and did not interact with either CO₂ or nutrient heterogeneity. Under elevated CO₂ conditions, the two- and three-species mixtures showed a clear trend towards underyielding. Our results show that differences among composition levels were dependent on soil heterogeneity, highlighting its potential role in modulating diversity–productivity relationships. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible to authorized users.     

Invading rainbow trout usurp a terrestrial prey subsidy from native charr and reduce their growth and abundance

Movements of prey organisms across ecosystem boundaries often subsidize consumer populations in adjacent habitats. Human disturbances such as habitat degradation or non-native species invasions may alter the characteristics or fate of these prey subsidies, but few studies have measured the direct effects of this disruption on the growth and local abundance of predators in recipient habitats. Here we present evidence, obtained from a combined experimental and comparative study in northern Japan, that an invading stream fish usurped the flux of allochthonous prey to a native fish, consequently altering the diet and reducing the growth and abundance of the native species. A large-scale field experiment showed that excluding terrestrial invertebrates that fell into the stream with a mesh greenhouse reduced terrestrial prey in diets of native Dolly Varden charr (Salvelinus malma) by 46–70%, and reduced their growth by 25% over six weeks. However, when nonnative rainbow trout (Oncorhynchus mykiss) were introduced, they monopolized these prey and caused an even greater reduction of terrestrial prey in charr diets of 82–93%, and reduced charr growth by 31% over the same period. Adding both greenhouse and rainbow trout treatments together produced similar results to adding either alone. Results from a comparative field study of six other stream sites in the region corroborated the experimental findings, showing that at invaded sites rainbow trout usurped the terrestrial prey subsidy, causing a more than 75% decrease in the biomass of terrestrial invertebrates in Dolly Varden diets and forcing them to shift their foraging to insects on the stream bottom. Moreover, at sites with even low densities of rainbow trout, biomass of Dolly Varden was more than 75% lower than at sites without rainbow trout. Disruption of resource fluxes between habitats may be a common, but unidentified, consequence of invasions, and an additional mechanism contributing to the loss of native species Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.