Resource supplements cause a change in colony sex-ratio specialization in the mound-building ant, Formica exsecta

We examine the role of food resources on split sex ratios in Formica exsecta. Models of resource-based sex allocation predict that greater resources will cause an increase in the production of reproductive females (gynes) and an increase in overall size of offspring. We experimentally increased food resources for a subset of colonies in a polygynous population with a very male-biased sex ratio. This increase in food availability caused colonies that were male specialists the prior year to switch to female production. Overall, a significantly greater proportion of food-supplemented colonies produced gynes, compared to control colonies. Moreover, food-supplemented colonies produced significantly larger workers and males (but not gynes), compared to those produced by control colonies. There was, however, no significant difference in the numerical productivity of food-supplemented and control colonies. We also measured the natural association between colony sex specialization and proximity to conifers, which typically harbor honeydew-bearing aphids (an important natural food source). In line with the view that resources play an important role for determining sex ratios in social insects, we found that female-producing colonies were significantly closer to conifers than were male-producing colonies

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 V. 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

Sex allocation in mound-building ants: The roles of resources and queen replenishment

Social Hymenoptera have become key organisms for tests of sex-ratio theory. We assess the role of resources for explaining sex-ratio variation in a highly male-biased population of the ant Formica exsecta. Key predictions of two of the three leading hypotheses invoking an effect of resource availability on sex ratios in social insects are not upheld. One prediction of the multifaceted parental-investment hypothesis is that colonies with greater brood production will rear a greater proportion of their diploid offspring as reproductive females (gynes), rather than workers, This study shows the positive correlation between female-biased sex ratio and the production of brood by female-producing colonies was not the outcome of a trade-off in the production of workers vs. gynes. The main prediction of the constant-female hypothesis is that investment in gynes should be constant. We found extreme variation among colonies in the number of gynes produced. By contrast, all the predictions of the queen-replenishment hypothesis were supported. The queen-replenishment hypothesis predicts that colonies produce gynes only when queen number is so low that colony production of brood is reduced, or colony survival threatened. We found that, as predicted by this hypothesis, female-producing colonies had fewer queens and produced a lower biomass of brood than male-producing colonies. In contrast, male-producing colonies had lower production per queen, and were more likely to be limited by external resources. This suggests that queen number limits production in female-producing colonies. These data provide evidence of adaptive adjustment of resources to worker, gyne, and male brood

Sex-ratio dependent execution of queens in polygynous colonies of the ant Formica exsecta

Formica exsecta has become an important model system for studying intraspecific variation in sex ratios. Patterns of sex allocation in polygynous (multiple queen per nest) populations of F. exsecta are generally consistent with the queen-replenishment hypothesis. This hypothesis states that colonies produce gynes (reproductive females) in order to increase queen number and enhance colony survival and/or productivity when the number of resident queens is low. However, the small proportion of colonies that raise gynes produce more than necessary for simple queen replenishment. It has been hypothesized that excess production of gynes may occur to reduce the frequency of accepting foreign unrelated gynes into the colony when workers cannot distinguish nestmate from non-nestmate queens. This explanation for excess gynes requires weak or no aggression between non-nestmates and is expected to lead to the selective execution of new queens by colonies that do not invest in the production of gynes. Experimental studies where gynes were introduced into natal and foreign colonies indeed suggested that polygynous populations of F. exsecta have a poor nestmate recognition system. Although gynes were significantly more likely to be accepted in their parental colony compared to another foreign female-producing colony, the difference was small. Moreover, encounters between workers from different colonies within the population showed very little aggression and were no more aggressive than encounters between nestmates, again suggesting a weak capacity for nestmate recognition. Our experiment also showed that colonies that produced only males executed most of the gynes that were experimentally introduced into the colony, whereas female-producing colonies accepted most gynes. This is consistent with ants using a simple rule of thumb to decrease parasitism by unrelated queens, whereby colonies selectively destroy gynes whenever gynes are not produced in the colonies

Temporal and spatial variations of gyne production in the ant Formica exsecta

Social insects have become a general model for tests of sex allocation theory. However, despite tremendous interest in the topic, we still know remarkably little about the factors that cause dramatic differences in sex allocation among local populations. A number of studies have suggested that environmental factors may influence sex allocation in ant populations. In polygynous (multiple queens per nest) populations of the ant Formica exsecta, sex allocation is extremely male biased at the population level, with only a small proportion of nests producing any gynes (female reproductive brood). We analysed the proportion of gyne-producing nests in 12 F. exsecta populations during three successive breeding seasons and found considerable temporal and spatial variability in the proportion of gyne-producing nests. The populations differed in a number of characteristics, including elevation, nest density, size of the nest mound, and number of nests per population. However, the proportion of gyne-producing nests was not associated with any of these geographic and demographic variables. Moreover, differences between populations in the production of gynes were not consistent between years. Thus, the proportion of gyne-producing nests appears to vary stochastically, perhaps because of stochastic variations in environmental factors. For example, year-to-year variations in the proportion of gyne-producing nests were associated with differences in spring weather conditions between years. The finding that gyne production varies greatly between years suggests that it may not always be adaptive at a local scale

(Contravariant) Koszul duality for DG algebras

A DG algebras $A$ over a field $k$ with $H(A)$ connected and $H_{<0}(A)=0$ has a unique up to isomorphism DG module $K$ with $H(K)\cong k$. It is proved that if $H(A)$ is degreewise finite, then RHom_A(?,K): D^{df}_{+}(A)^{op} \equiv D_{df}^{+}}(RHom_A(K,K)) is an exact equivalence of derived categories of DG modules with degreewise finite-dimensional homology. It induces an equivalences of $D^{df}_{b}(A)^{op}$ and the category of perfect DG $RHom_A(K,K)$-modules, and vice-versa. Corresponding statements are proved also when $H(A)$ is simply connected and $H^{<0}(A)=0$.Comment: 33 page

Spectroscopy of Bright QUEST RR Lyrae Stars: Velocity Substructures toward Virgo

Using a sample of 43 bright (V<16.1, distance <13 kpc) RR Lyrae stars (RRLS) from the QUEST survey with spectroscopic radial velocities and metallicities, we find that several separate halo substructures contribute to the Virgo overdensity (VOD). While there is little evidence for halo substructure in the spatial distribution of these stars, their distribution in radial velocity reveals two moving groups. These results are reinforced when the sample is combined with a sample of blue horizontal branch stars that were identified in the SDSS, and the combined sample provides evidence for one additional moving group. These groups correspond to peaks in the radial velocity distribution of a sample of F type main-sequence stars that was recently observed in the same directon by SEGUE, although in one case the RRLS and F star groups may not lie at the same distance. One of the new substructures has a very narrow range in metallicity, which is more consistent with it being the debris from a destroyed globular cluster than from a dwarf galaxy. A small concentration of stars have radial velocities that are similar to the Virgo Stellar Stream (VSS) that was identified previously in a fainter sample of RRLS. Our results suggest that this feature extends to distances as short as ~12 kpc from its previous detection at ~19 kpc. None of the new groups and only one star in the sample have velocities that are consistent with membership in the leading tidal stream from the Sagittarius Dwarf Spheroidal Galaxy, which some authors have suggested is the origin of the VOD.Comment: Accepted for publication in the A

Probing Solar Convection

In the solar convection zone acoustic waves are scattered by turbulent sound speed fluctuations. In this paper the scattering of waves by convective cells is treated using Rytov's technique. Particular care is taken to include diffraction effects which are important especially for high-degree modes that are confined to the surface layers of the Sun. The scattering leads to damping of the waves and causes a phase shift. Damping manifests itself in the width of the spectral peak of p-mode eigenfrequencies. The contribution of scattering to the line widths is estimated and the sensitivity of the results on the assumed spectrum of the turbulence is studied. Finally the theoretical predictions are compared with recently measured line widths of high-degree modes.Comment: 26 pages, 7 figures, accepted by MNRA

Patterns of Natural and Human-Caused Mortality Factors of a Rare Forest Carnivore, the Fisher (Pekania pennanti) in California.

Wildlife populations of conservation concern are limited in distribution, population size and persistence by various factors, including mortality. The fisher (Pekania pennanti), a North American mid-sized carnivore whose range in the western Pacific United States has retracted considerably in the past century, was proposed for threatened status protection in late 2014 under the United States Endangered Species Act by the United States Fish and Wildlife Service in its West Coast Distinct Population Segment. We investigated mortality in 167 fishers from two genetically and geographically distinct sub-populations in California within this West Coast Distinct Population Segment using a combination of gross necropsy, histology, toxicology and molecular methods. Overall, predation (70%), natural disease (16%), toxicant poisoning (10%) and, less commonly, vehicular strike (2%) and other anthropogenic causes (2%) were causes of mortality observed. We documented both an increase in mortality to (57% increase) and exposure (6%) from pesticides in fishers in just the past three years, highlighting further that toxicants from marijuana cultivation still pose a threat. Additionally, exposure to multiple rodenticides significantly increased the likelihood of mortality from rodenticide poisoning. Poisoning was significantly more common in male than female fishers and was 7 times more likely than disease to kill males. Based on necropsy findings, suspected causes of mortality based on field evidence alone tended to underestimate the frequency of disease-related mortalities. This study is the first comprehensive investigation of mortality causes of fishers and provides essential information to assist in the conservation of this species