Chemical communication of queen supergene status in an ant

Traits of interest to evolutionary biologists often have complex genetic architectures, the nature of which can confound traditional experimental study at single levels of analysis. In the fire ant Solenopsis invicta, the presence of a Mendelian ‘supergene’ is both necessary and sufficient to induce a shift in a fundamental property of social organization, from single‐queen (monogyne) to multiple‐queen (polygyne) colonies. This selfish genetic element, termed the Social b (Sb) supergene, contains > 600 genes that collectively promote its fitness by inducing the characteristic polygyne syndrome, in part by causing polygyne workers to accept only queens bearing the Sb element (a behaviour termed ‘worker Sb discrimination’). Here, we employ a newly developed behavioural assay to reveal that polygyne workers, many of which bear the Sb element, employ chemical cues on the cuticle of queens to achieve worker Sb discrimination, but we found no evidence for such pheromonally mediated worker Sb discrimination in monogyne workers, which universally lack the Sb element. This polygyne worker Sb discrimination was then verified through a ‘green beard’ effect previously described in this system. We thus have demonstrated that the Sb element is required both for production of relevant chemical cues of queens and for expression of the behaviours of workers that collectively result in worker Sb discrimination. This information fills a critical gap in the map between genotype and complex phenotype in S. invicta by restricting the search for candidate genes and molecules involved in producing this complex social trait to factors associated with the Sb element itself.     

Carbon isotope discrimination in leaves of the broad‐leaved paperbark tree,Melaleuca quinquenervia,as a tool for quantifying past tropical and subtropical rainfall

Quantitative reconstructions of terrestrial climate are highly sought after but rare, particularly in Australia. Carbon isotope discrimination in plant leaves (Δ_leaf) is an established indicator of past hydroclimate because the fractionation of carbon isotopes during photosynthesis is strongly influenced by water stress. Leaves of the evergreen tree Melaleuca quinquenervia have been recovered from the sediments of some perched lakes on North Stradbroke and Fraser Islands, south‐east Queensland, eastern Australia. Here, we examine the potential for using M. quinquenervia ?_leaf as a tracer of past rainfall by analysing carbon isotope ratios (δ¹³C) of modern leaves. We firstly assess Δ_leaf variation at the leaf and stand scale and find no systematic pattern within leaves or between leaves due to their position on the tree. We then examine the relationships between climate and Δ_leaf for a 11‐year time series of leaves collected in a litter tray. M. quinquenervia retains its leaves for 1–4 years; thus, cumulative average climate data are used. There is a significant relationship between annual mean ?_leaf and mean annual rainfall of the hydrological year for 1–4 years (i.e. 365–1460 days) prior to leaf fall (r² = 0.64, P = 0.003, n = 11). This relationship is marginally improved by accounting for the effect of pCO₂ on discrimination (r² = 0.67, P = 0.002, n = 11). The correlation between rainfall and Δ_leaf, and the natural distribution of Melaleuca quinquenervia around wetlands of eastern Australia, Papua New Guinea and New Caledonia offers significant potential to infer past rainfall on a wide range of spatial and temporal scales.     

Assessing the effects of sodium on fire ant foraging in the field and colony growth in the laboratory

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Predicting socio‐economic and biodiversity impacts of invasive species: Red Imported Fire Ant in the developing western Pacific

Effective biosecurity and pest management are fundamental to sustainable development. Invasive ants pose significant risks to the environment and economy, which are well‐managed by biosecurity agencies in developed countries. However, a comprehensive view of the potential impacts of invasive ants in developing Pacific Island Countries and Territories (PICTs) is lacking. We quantified the potential costs of the Red Imported Fire Ant (Solenopsis invicta Buren) across multiple sectors using an extrapolation analysis. Overall, we estimated that the impacts of Red Imported Fire Ant on developing PICTs could amount to over USD 329 million annually, corresponding to approximately 0.7% of combined GDP. Over half of the costs were predicted to result from impacts on the agriculture sector, a major source of employment and subsistence. We found that over 350 highly threatened species could be at risk from Red Imported Fire Ant. We would expect countries with Least Developed Country status and relatively low GDP to be least able to respond to an incursion of these ants, and as a result the costs could be higher than we have extrapolated. Red Imported Fire Ant could therefore potentially have considerable impact on the on‐going development of the region.     

Population recovery and spatial dynamics of colony recruitment in a harvester ant: a case study in a park

The recovery process of Messor aciculatus (Fr. Smith) harvester ant colonies from habitat disturbance by construction work in an urban park was investigated from 1989 to 1994. Colonies located in 1989 were considered to be survivors of the disturbance, and colonies located for the first time after 1990 were considered to be recruits. The colonies continuously increased in number after the disturbance, and the population size more than doubled just after the disturbance. Analyses of the nest site distribution of colonies in 1989 and 1994 suggest that landform modification destroyed some of them, and that the changed nest site distribution in 1989 affected the distribution in 1994, or, in other words, the disturbance indirectly affected the nest distribution 5 years later. Analyses of the nearest neighbor to each colony suggested that established colonies would obstruct newly recruited colonies around their nests. Construction may open up space for recruitment colonization by removing established colonies, and may also create suitable conditions by destruction of original vegetation, or returning vegetation to the early stages of plant community succession. As a result, the disturbance may facilitate the subsequent recruitment of new colonies and rejuvenate the population.     

Effect of park age,size, shape and isolation on ant assemblages in two cities of Southern Spain

In the context of conservation biology, we have searched for the most influential characteristics of urban parks on ecological diversity. In this study, therefore, we studied the ant community of 24 parks in two cities of Southern Spain (Cordoba and Seville). In total, 46 species were found. Abundant and widely distributed species were synanthropic ones, such as Lasius grandis, Pheidole pallidula or Tapinoma nigerrimum, and exotic species such as Cardiocondyla mauritanica. However some parks also harbored natural habitat‐preferring species. Park size, shape and distance to city outskirts were the most important predictors of species richness and diversity. Likewise, park age influenced the composition of the ant fauna.     

Stable isotope analysis and fin whale subpopulation structure in the eastern North Atlantic

The North Atlantic fin whale was subject to heavy exploitation in the past and, despite partial recovery, it is still considered endangered. Recent research has questioned its currently accepted subpopulation structure and migratory movements, challenging management and conservation efforts. Here we contribute to this discussion by presenting results of stable isotope analysis of fin whale bones and krill samples collected from fin whale stomachs from two exploited areas, west Iceland and northwest Spain, and comparing these results with North Atlantic isoscapes. In krill, δ¹⁵N values were highly variable and no significant differences in δ¹⁵N and δ¹³C between areas emerged. δ¹⁵N and δ¹³C values in bone were higher than in krill, due to trophic enrichment, and were not significantly different between areas. Both krill and bone results were slightly inconsistent with the local isoscapes, maybe due to seasonal variations in local productivity and in krill diet and, in the case of bone, to its capability to integrate long‐term isotopic values, derived from food consumed in distant areas. Conversely, δ¹⁸O values, which reflect the basal signal of sea water, were consistent with isoscapes and significantly higher in whales from northwestern Spain, possibly indicating migration to breeding grounds located at lower latitudes.     

The time scale of isotope signals in spiders: molting the remains of a previous diet

Stable isotope analysis (SIA) has emerged as an important tool for understanding consumer diets and diet shifts. However, although the general idea behind SIA is clear, the interpretation of data is often fraught with problems because tissue turnover and fractionations are not known. We investigated shifts in stable isotope composition of spiders following a diet shift, using mealworms fed either maize (C4) or wheat (C3) flour. Mealworms had different carbon isotope composition depending on their diet and this difference was reflected in spider body parts. In the experiment, we first fed the spiders on a diet of either maize‐fed or wheat‐fed mealworms and then switched diet at the time of the second molt. Spiders were then sampled repeatedly until the next molt. We sampled both legs and abdomens, as these are presumed to have different turnover of tissue, and also molt remains were sampled when this was relevant. The data indicated that the spider legs had a turnover of about 20 days, whereas the spider abdomens had a turnover of about 8 days. Molt remains had the slowest turnover and reflected the diet at the previous molt, when the exoskeleton was formed. Both these observations indicate that SIA may be successfully used for elucidating diet shifts. More problematic was the fact that fractionation of carbon isotope ratios varied with body parts and diets. When spiders were fed maize‐mealworms then the fractionation was larger for abdomens, but when the spiders were fed wheat‐mealworms then the fractionation was larger for legs. The mechanisms underlying this pattern are unclear and deserve further attention.     

Stable isotope records of sei whale baleens from Chilean Patagonia as archives for feeding and migration behavior

Carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope variations in baleen plates of sei whales (Balaenoptera borealis) stranded after a mass mortality event in Chilean Patagonia were investigated to assess potential dietary and migratory patterns. Carbon and nitrogen isotope ratios of seven baleens from six individuals were analyzed. The δ¹³C values ranged from ? 19.1 to ? 15.9‰ and the δ¹⁵N values from 8.7 to 15.4‰. Variations of up to 2.9‰ for δ¹³C and 5.3‰ for δ¹⁵N were observed within one baleen. Carbon and nitrogen isotope records of each baleen were significantly correlated and showed recurring oscillations confirmed by wavelet analyses. Oscillations slightly differed in periodicity indicating variable baleen growth rates between 10.0 and 16.5 cm/year. Food sources of the whales are discussed in terms of available isotope data for potential prey taxa and potential migratory behavior on the basis of latitudinal isotope gradients of particulate organic matter. Cyclicity could be explained by regular migrations of the sei whales from subtropical calving areas to high‐latitude foraging grounds. δ¹⁵N records of baleens differed between individuals eventually pointing to diverse feeding and migratory preferences among sei whale individuals.     

Genotypic variations in carbon isotope discrimination, transpiration efficiency, and biomass production in rice as affected by soil water conditions and N

Genotypic and environmental (soil water regime and N level) variation in carbon isotope discrimination (CID) in relation to the gas exchange, transpiration efficiency (A/T), and biomass production were investigated in field experiments using eleven rice (Oryza sativa L.) genotypes. The results showed that genotype was more dominant for variation in CID than in total biomass. Genotypic ranking in CID was consistent across environments because of small genotype × environment interactions. Japonica genotypes tended to have lower CID than indica genotypes. Higher soil water and lower N rate significantly increased CID. Variation in CID was slightly smaller for water regime than for genotype. There was a negative correlation between CID andA/T among genotypes within water regimes. Genotypic variation in CID was associated mainly with variation in stomatal conductance under all soil water regimes and with photosynthetic capacity in late growth stages under aerobic soil conditions. The decrease in CID at higher N was probably due to lower stomatal conductance under aerobic soil conditions and to higher photosynthetic rates under submerged soil conditions. The correlation between biomass and CID was not clear in aerobic soil, whereas it was positive in submerged soil, which indicated that the significance of lower or higher CID for improving biomass productivity may differ under different soil water regimes. Overall, the results implied a possible use of CID as a selection criterion for genotypic improvement inA/T and productivity in rice.     

Diet of harbor porpoises along the Dutch coast: A combined stable isotope and stomach contents approach

High stranding frequency of porpoises, Phocoena phocoena, along the Dutch coast since 2006 has led to increased interest in the ecology of porpoises in the North Sea. Stranded porpoises were collected along the Dutch coast (2006–2008) and their diet was assessed through stomach content and stable isotope analysis (δ¹³C and δ¹⁵N) of porpoise muscle and prey. Stable isotope analysis (SIAR) was used to estimate the contribution of prey species to the porpoises' diet. This was compared to prey composition from stomach contents, to analyze differences between long‐ and short‐term diet. According to stomach contents, 90.5% of the diet consisted of gobies, whiting, lesser sandeel, herring, cod, and sprat. Stable isotope analysis revealed that 70‐83% of the diet consisted of poor cod, mackerel, greater sandeel, lesser sandeel, sprat, and gobies, highlighting a higher importance of pelagic, schooling species in the porpoises' diet compared to stomach contents. This could be due to prey distribution as well as differences in behavior of porpoises and prey between the coastal zone and offshore waters. This study supports the need for multi‐method approaches. Future ecological and fishery impact assessment studies and management decisions for porpoise conservation should acknowledge this difference between the long‐ and short‐term diet.     

Stable isotope evidence for sex- and status-based variations in diet and life history at medieval Trino Vercellese, Italy

The medieval period in Europe was a time of unprecedented social complexity that affected human diet. The diets of certain subgroups-for example, children, women, and the poor-are chronically underrepresented in historical sources from the medieval period. To better understand diet and the distribution of foods during the medieval period, we investigated stable carbon and nitrogen isotope ratios of 30 individuals from Trino Vercellese, Northern Italy (8th-13th c.). Specifically, we examined diet differences between subgroups (males and females, and high- and low-status individuals), and diet change throughout the life course among these groups by comparing dentine and bone collagen. Our results show a diet based on terrestrial resources with input from C(4) plants, which could include proso and/or foxtail millet. Diets of low-status males differ from those of females (both status groups) and of high-status males. These differences develop in adulthood. Childhood diets are similar among the subgroups, but sex- and status-based differences appear in adulthood. We discuss the possibility of cultural buffering and dietary selectivity of females and high-status individuals.     

The roles of foraging environment,host species,and host diet for a generalist pupal parasitoid

Even for parasitoids with a wide host range, not all host species are equally suitable, and host quality often depends on the plant the host feeds on. We compared oviposition choice and offspring performance of a generalist pupal parasitoid, Pteromalus apum (Retzius) (Hymenoptera: Pteromalidae), on two congeneric hosts reared on two plant species under field and laboratory conditions. The plants contain defensive iridoid glycosides that are sequestered by the hosts. Sequestration at the pupal stage differed little between host species and, although the concentrations of iridoid glycosides in the two plant species differ, there was no effect of diet on the sequestration by host pupae. The rate of successful parasitism differed between host species, depending on the conditions they were presented in. In the field, where plant‐associated cues are present, the parasitoid used Melitaea cinxia (L.) over Melitaea athalia (Rottemburg) (Lepidoptera: Nymphalidae), whereas more M. athalia were parasitised in simplified laboratory conditions. In the field, brood size, which is partially determined by rate of superparasitism, depended on both host and plant species. There was little variation in other aspects of offspring performance related to host or plant species, indicating that the two host plants are of equal quality for the hosts, and the hosts are of equal quality for the parasitoids. Corresponding to this, we found no evidence for associative learning by the parasitoid based on their natal host, so with respect to these host species they are truly generalist in their foraging behaviour.     

Fine‐scale variation in natural nitrogen isotope ratios of ants

Stable isotopes provide a powerful means of elucidating the trophic ecology of organisms. Analyses of variation in the ratio of nitrogen isotopes (δ¹⁵N) can provide insights into the trophic position of species with broad diets and the ability to occupy multiple positions in food webs, such as ants. The most powerful studies compare subjects across various spatial scales, but to do so, local variation in δ¹⁵N baselines must be taken into account. To date, a wide variety of baseline calibration methods have been employed, leading some authors to suggest that a standard approach is needed, and that the reality of environmental variation necessitates that this should be at fine scales. In this study, we examine the fine‐scale variation in δ¹⁵N value of colonies of the ant Formica kozlovi Dlussky (Hymenoptera: Formicidae: Formicini) along a sloped transect in Mongolia, and compare these with values for associated soils in an effort to shed further light on this issue. We find variation in ant δ¹⁵N to the order of one trophic level (ca. 3‰), over a distance of only 1 km. Ant δ¹⁵N was highly correlated with soil δ¹⁵N, and variation in mineral soil δ¹⁵N explained ca. 81% of the variation in ant δ¹⁵N. This study underlines the importance of local‐scale baseline corrections for isotopic studies, particularly in environments where baseline variation might be expected. It further suggests that δ¹⁵N of mineral soils may provide a suitable baseline for ecological studies of terrestrial arthropods.     

Interacting effects of atmospheric CO2 enrichment and solar radiation on growth of the aquatic fern Azolla filiculoides

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Effects of land‐use changes on ecosystem services: decrease in ant predation in human‐dominated landscapes in central Brazil

Anthropogenic disturbances often affect the abundance and diversity of ants (Hymenoptera: Formicidae) but relatively few studies have explored the implications of such changes on the ecosystem services mediated by these insects. Here, we evaluated how the transformation of Cerrado savanna habitats into crop plantations affects the abundance, diversity, and the predatory activity of ants. A survey of the ant faunas foraging above‐ and belowground was performed in six crop and six non‐crop (i.e., native vegetation) sites. Above‐ and belowground rates of ant predation were estimated at these same sites using mealworms, Tenebrio molitor L. (Coleoptera: Tenebrionidae), as baits, simulating crop herbivores. Belowground predation rates were significantly greater in the non‐crop sites, despite the lack of difference in overall abundance and species richness of ants foraging belowground between the crop vs. non‐crop sites. In contrast, we did not detect any significant difference in aboveground predation rates between crop vs. non‐crop sites even though there were significantly more species of ants foraging aboveground in the non‐crop sites. Army ants (subfamily Dorylinae) were the main predatory species belowground, and their abundance was significantly greater in non‐crop sites. In contrast, the main predators aboveground were omnivore ants of the genera Pheidole and Solenopsis, which had similar abundances in the crop and non‐crop sites. Overall, our results indicate that transformation of native Cerrado habitats into crop plantations reduces the abundance of some important predatory species, notably those that forage belowground, and this may negatively affect the potential for ants to provide pest control services in agroecosystems.     

The carbon fertilization effect over a century of anthropogenic CO2 emissions: higher intracellular CO2 and more drought resistance among invasive and native grass species contrasts with increased water use efficiency for woody plants in the US Southwest

From 1890 to 2015, anthropogenic carbon dioxide emissions have increased atmospheric CO₂ concentrations from 270 to 400 mol mol^?1. The effect of increased carbon emissions on plant growth and reproduction has been the subject of study of free‐air CO₂ enrichment (FACE) experiments. These experiments have found (i) an increase in internal CO₂ partial pressure (c_i) alongside acclimation of photosynthetic capacity, (ii) variable decreases in stomatal conductance, and (iii) that increases in yield do not increase commensurate with CO₂ concentrations. Our data set, which includes a 115‐year‐long selection of grasses collected in New Mexico since 1892, is consistent with an increased c_i as a response to historical CO₂ increase in the atmosphere, with invasive species showing the largest increase. Comparison with Palmer Drought Sensitivity Index (PDSI) for New Mexico indicates a moderate correlation with Δ¹³C (r² = 0.32, P < 0.01) before 1950, with no correlation (r² = 0.00, P = 0.91) after 1950. These results indicate that increased c_i may have conferred some drought resistance to these grasses through increased availability of CO₂ in the event of reduced stomatal conductance in response to short‐term water shortage. Comparison with C₃ trees from arid environments (Pinus longaeva and Pinus edulis in the US Southwest) as well as from wetter environments (Bromus and Poa grasses in New Mexico) suggests differing responses based on environment; arid environments in New Mexico see increased intrinsic water use efficiency (WUE) in response to historic elevated CO₂ while wetter environments see increased c_i. This study suggests that (i) the observed increases in c_i in FACE experiments are consistent with historical CO₂ increases and (ii) the CO₂ increase influences plant sensitivity to water shortage, through either increased WUE or c_i in arid and wet environments, respectively.