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The dominance-nutrition hypothesis predicts that nutritional intake and energetic costs in adulthood interact to drive behavioral and physiological differences between females in primitively eusocial insects, and thereby affect reproductive caste. We tested predictions of this hypothesis in independent-founding Mischocyttarus pallidipectus paper wasps. We measured stable isotope tissue composition to compare nutritional status before adult emergence and among adult females. Adult tissue δ15N content (an indicator of feeding at a higher trophic level because it is enriched in animal prey relative to plant-based foods) was significantly higher in adults than at the end of pupal development, suggesting adult nutrition affects δ15N content after the end of pupal development. We then asked if behavior and nutritional status predicted ovary development. We measured ovary development, nitrogen and carbon stable isotope ratios, dominance behavior, and task performance (foraging, as indicated by time spent on the nest) for adult female wasps. We used social network analysis to quantify differences in social status between females with developed and undeveloped (filamentous) ovaries. Dominant females spent more time on the nest and were significantly more enriched in δ15N than subordinate females. These data support the dominance-nutrition hypothesis: adult behavior and energy expenditure, and access to animal-based diets, correspond to female reproductive physiology, and may play a role in adult-stage caste determination. Within animal social groups, differential access to resources can affect differences in reproductive success. This may affect caste determination in eusocial colonies, where reproductive tasks are divided between female castes, with a queen (or multiple queens) capable of egg-laying and helped by sterile workers. Female sterile workers and reproductives of some social Hymenoptera are morphologically similar at adult emergence, suggesting that adult experience may impact caste status. We asked whether patterns of nutritional physiology in Mischocyttarus pallidipectus paper wasps matched adult-stage caste determination by testing predictions of the dominance-nutrition hypothesis, which states that adult nutrition and energetic costs of tasks shape reproductive female development through behavioral and physiological changes.

When helpers from cooperative breeding animals have some expectation of direct reproduction, there is potential for conflict over how much aid they should provide to the colony. For example, if food is shared among all colony members, then higher levels of foraging by a helper would be desirable for the colony as a whole. However, because foraging is risky and physiologically costly, hopeful reproductive helpers could avoid foraging. Evidence suggests that this work-conflict could be resolved if helpers are aggressively coerced by their nestmates to provide aid. Here, we showed that in the primitively eusocial paper wasp Polistes versicolor, colony food starvation leads to an increasing in aggression that results in an increasing activity level (including foraging). We propose that aggression affects forage levels because (i) attacks from nestmates are directed toward known foragers rather than non-foragers; and (ii) resting wasps generally respond to aggression by becoming more active while already active wasps generally respond by switching the task they were doing. In P. versicolor, direct reproduction is an option for helpers. It means that they can be considered as hopeful reproductive individuals seeking to avoid performing risky behaviours, like foraging. In this sense, decentralized aggression from nestmates could be a coercive mechanism to force wasps performing undesirable tasks, while simultaneously enhancing the performance of a variety of other tasks.

In cooperative breeders, subordinates that have alternative reproductive options are expected to stay and help dominant breeders only as long as they contribute to group productivity, if their fitness is linked with colony success. Female Polistes dominula paper wasps live as cooperative breeders in small groups of typically fewer than 10 females. Subordinates tend to have high-quality outside options, and so could choose alternative breeding tactics if their work efforts increased productivity negligibly. In the founding stage before workers emerge, we tested the effect of various predictors on nest growth, as a proxy for group productivity, and explored the shape of the relationship between group size and nest growth. We found group size to be the only significant predictor of nest growth: variation among body sizes within the group showed no effect, suggesting a lack of size-dependent task specialization in this species. Average body size and average genetic relatedness between group members similarly showed no effects on nest growth. Group size had a non-linear effect so that per-capita benefits to nest growth decreased in larger groups, and groups of 10 or more would benefit negligibly from additional group members. Hence, females might be better off pursuing other options than joining a large group. This finding helps to explain why P. dominula groups are usually relatively small in our study population. Further studies may illuminate the mechanisms behind the smaller per-capita nest growth that we found in larger groups. Significance statement Identifying which factors influence the productivity of animal groups is key to understanding why different species breed cooperatively in groups of varying sizes. In the paper wasp Polistes dominula, we investigated the growth rate of nests as a measure of group productivity. We found that average body size, the variation in body sizes within the group, and average genetic relatedness between group members did not affect nest growth, while group size had a strong, positive effect: nests grew faster with more group members, but the per-capita benefit decreased in larger groups. The addition of extra group members in groups of 10 or more had negligible effects on nest growth. Hence, wasps may be better off pursuing other options than joining large groups. This finding helps to explain why groups normally consist of fewer than 10 wasps in this population. Electronic supplementary material The online version of this article (10.1007/s00265-018-2502-x) contains supplementary material, which is available to authorized users.

Division of labor in social groups can be influenced by differential nutrition. Consumption of more food or higher-quality food often affects individuals’ capacities for reproduction. In social insects, nutrients consumed during immature (e.g., larval) stages often affect adult reproductive capacity, but adult nutrition may also impact reproductive status. This study tested whether ovary development, an indicator of reproductive status, corresponded to higher trophic-level feeding for adults in the primitively eusocial paper wasp Mischocyttarus mastigophorus. Our main prediction was that females’ ovary development would correlate positively with evidence of feeding at higher trophic levels, as indicated by stable isotope ratios of nitrogen (δ15N) and carbon (δ13C). We first asked whether isotope ratios of mature females co-varied with ovary development. δ15N values were higher for mature females with better-developed ovaries, as expected if they fed on a diet richer in animal tissue. There was a negative relationship of δ13C with ovary development in mature females, as would be expected if females with developed ovaries had higher body lipid stores. To test for evidence of nutritional biasing of caste during immature development, we measured changes in isotope ratios across pupal development leading up to early adulthood (i.e., immediately before and after adult eclosion). The δ15N and δ13C values for mature pupae were similar to those of newly emerged adults and to those of mature adults lacking developed ovaries. In contrast, mature females with developed ovaries showed N-isotope signatures of a more prey-based diet and C-isotope signatures of elevated lipid content. We conclude the N and C isotopic signatures of ovary-developed mature females diverged from their levels at the end of immature development. The findings suggest reproductive caste status was associated with differences in nutrient acquisition and reflects differences in consumption of animal-derived versus plant-based foods during adulthood. Pre-adult nutrition is known to affect caste development, and thereby influence division of labor, in diverse insect societies. In temperate independent-founding (IF) paper wasps, the effects of larval nutrition on adult caste may reflect adaptations for overwintering and diapause by reproductive females. We asked whether adult nutritional content showed caste-related trophic differences in the Neotropical IF paper wasp Mischocyttarus mastigophorus. Previous studies suggested interactions among adult nest-mates affected food sharing and caused biased nutrient flow within colonies. Our data suggest adult nutrition affected reproductive physiology. We propose this pattern may be widespread in tropical IF paper wasps, where opportunities for adult reproductive plasticity are favored by long colony cycles and relatively aseasonal nest founding.

Inter-specific variation in the occurrence of individual recognition (IR) has mostly been examined in species that differ with respect to two critical conditions necessary for such ability: presence of complex social behaviours and sender’s distinctiveness. However, variation in IR when species share such conditions is lesser known. We tested for IR in the paper wasp Polistes versicolor and compared the results with published information on IR in Polistes fuscatus. Both species have a flexible nest-founding behaviour and form dominance hierarchies. Additionally, both have individually distinctive facial patterns. P. versicolor faces are highly variable, and they do not appear to be associated with individual quality, consistent with the role as cues in IR. However, the wasps did not discriminate between nestmates with familiar and unfamiliar faces. Besides, wasps did not discriminate between familiar and unfamiliar social partners. Familiar discrimination (discrimination of individuals based on prior social interactions) is considered the first stage toward IR; therefore, P. versicolor wasps, unlike P. fuscatus, lack the ability for IR. Our results suggest that the high variation in colour pattern may not be a good proxy for the occurrence of IR in paper wasps and that neutral variation may be an underestimated factor promoting phenotypic variability. High inter-individual phenotypic variability in primitively eusocial paper wasps has sometimes been considered to be a result of selection for efficient individual identity signalling, especially because senders may benefit from being individually recognised. P. versicolor paper wasp females have highly variable inter-individual facial patterns. However, we demonstrated that they lack individual recognition. After excluding some potentially confounding hypothesis, we conclude that selection for efficient individual identity signalling is not the only factor promoting phenotypic variability in paper wasps and that at least in some species, neutral variation may play a role.

We investigated the effect of kinship on mate selection by males of the neotropical primitively eusocial paper wasp Polistes versicolor. By conducting short-time, paired, consecutive, dyadic encounters, in which a male was randomly introduced separately to a related and an unrelated female in a small glass arena, we were able to observe and measure the male sexual display. We found that male sexual interest toward a female was not affected by kinship, even after controlling the differences in female behavior. Despite our finding that males were as eager to harass females, regardless of kinship, previous evidence suggested that incestuous copulation (inbreeding) in natural populations of P. versicolor appeared to be avoided. We suggest that in the natural mating habitat of the native population that we studied, in which many colonies were able to produce males and reproductive females in a short time window, the probability of finding related sexual partners was low, hence, intersexual kin discrimination did not evolve.

Various animals build nests with defensive structures to deter predation on offspring. Construction of nest defensive structures can reduce the probability of predation but will involve various costs. Here, we examined both the costs and benefits of the construction of a nest defensive structure in a paper wasp, Polistes chinensis antennalis, and clarify whether the paper wasp changes the level of defensive structures of nests depending on predation risk. A foundress (queen) of the paper wasp starts a colony in spring and maintains her nest alone until the emergence of workers. At this stage, pupae in the nests are sometimes preyed on by conspecifics of other nests. The intruder needs to break the cocoon, which seals the entrance of the cell, to extract the pupa from the cell. Foundresses often apply nest material (pulp) to the surface of cocoons in their nests. We found that pulp on a cocoon increased the time an intruder required to break the cocoon. This result shows that the pulp structure on cocoons helps to prevent predation on pupae. On the other hand, pulp on cocoons involved costs, including time required to collect pulp and being a potential obstacle to the emergence of workers from the cocoon. Additionally, we found that the amount of pulp on cocoons was greater in nests under higher predation risk than nests under lower predation risk. These results suggest that pulp on cocoons is a nest defensive structure, and foundresses adjusted the construction of the defensive structure depending on predation risk.

Cuticular hydrocarbons (CHCs) are information-rich signals in social insects that coordinate behaviors within nests. However, in some taxa, the precise information conveyed by CHCs is poorly understood. In particular, there is a debate over whether CHCs convey information about their bearer's dominance or fertility. Distinguishing between dominance and fertility signaling is difficult because fertility and rank are frequently correlated within social insect colonies. This study disentangles those relationships by examining CHCs of Polistes dominulus paper wasps during the early nest-founding stage before dominance and fertility become correlated. First, we confirm that dominance and fertility are not associated in early spring foundresses. Then we show that CHCs are more strongly associated with fertility than dominance. There was no relationship between cuticular hydrocarbon profiles and a wasp’s ability to dominate rivals through aggression, suggesting that CHCs are unlikely to provide information about dominance. However, there was a significant correlation between ovarian development and the CHC profile, suggesting that CHCs could convey valuable information about their bearer's fertility. Furthermore, our data provide a potential mechanism for chemical signaling of fecundity, as there is a relationship between endogenous juvenile hormone titer (a gonadotropin), degree of ovarian development, and the CHC profile. Hormonal regulation of CHC profile expression offers a physiological mechanism to coordinate behavior, physical state, and signal expression.

Female Polistes paper wasps are capable of independent nesting, yet many populations demonstrate a mixture of solitary and cooperative nest foundation. Previous studies of Polistes have found survival and/or productivity advantages of cooperative nest foundation compared to solitary nesting, and reproductive skew models have been designed to predict the dynamics of such flexible cooperation. In this paper, we examine the success of different nesting strategies in a previously unstudied population of Polistes aurifer in southern California. The colony cycle of this population is less synchronous than that of other temperate species, and the frequency of solitary nesting averages 86.2%. Our data suggest that this low rate of cooperative nest founding is adaptive, as demonstrated by the lack of survival or productivity advantages for cooperative foundress associations. Due to foundress turnover and nest foundation later in the season, many nests produce only one set of offspring. This results in a loss of the eusocial nature of some nests in the population. Data from a small sample of multifoundress nests show significant positive reproductive skew, despite concession model predictions that skew should be low in populations with low ecological constraints on independent nesting. This lack of support for the concessions skew model reflects a diminished incentive for cooperation.

Made available in DSpace on 2018-11-26T17:06:24Z (GMT). No. of bitstreams: 0 Previous issue date: 2016-08-01 Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) Based on species endemism, three biodiversity centers, called Ecological Corridors have been proposed as one of the main conservation strategies for the Atlantic Rain Forest. This study tested whether the organization of the social paper wasp assemblage fits those centers. A standardized protocol was used for sampling the social paper wasp fauna. The structural organization was estimated by Nonmetric Multidimensional Scaling (NMDS) based on the similarity indexes of Sorensen (qualitative data) and Morisita-Horn (quantitative data). Regressive models were applied to the first axes' site scores of the NMDS, to the latitudinal and altitudinal variations, and to the speciation and immigration probabilities predicted by the neutral theory for a metacommunity. Our results indicated that the social paper wasp assemblage is organized in a continuum, with two distinct biodiversity centers. The organization of the assemblage along the gradient was dependent on latitudinal and altitudinal variations and their interactions, and also on the speciation and immigration probabilities. Several studies have demonstrated that the current biodiversity patterns of the Atlantic Forest might be explained by the past climate and, consequently, by the connection between the Amazon and the Atlantic Forest. In addition, speciation and immigration probabilities strongly influence the compositional and structural variations of the social paper wasp assemblage along the latitudinal gradient. Univ Estadual Paulista, Inst Biociencias Letras & Ciencias Exatas, Dept Zool & Bot, Rua Cristovao Colombo 2265, BR-15054000 Sao Jose Do Rio Preto, SP, Brazil Univ Sao Paulo, Museu Zool, Ave Nazare 481, BR-04263000 Sao Paulo, SP, Brazil Univ Estadual Paulista, Inst Biociencias Letras & Ciencias Exatas, Dept Zool & Bot, Rua Cristovao Colombo 2265, BR-15054000 Sao Jose Do Rio Preto, SP, Brazil FAPESP: 98/05083-0 FAPESP: 01/08060-5

A recent study (Izzo et al., Behav Ecol Sociobiol 64: 857–864, 2010) reported that cuticular hydrocarbons (CHCs) correlate with fertility, not dominance, in the paper wasp Polistes dominulus thus contradicting the results of recent investigations which concluded that social dominance is the main determinant for CHC signatures in this species. We suggest here that different forms of dominance in the pre-nesting and post-nesting phases caused the apparently contradictory results. Thus the assumption that dominance behaviour in the pre-nesting stage is synonymous with dominance after colony foundation is incorrect. We provide standardised definitions for forms of “dominance” observed in the P. dominulus life cycle to avoid apparent discrepancies in the future among studies dealing with the same topics in different annual stages.

Successful Polistes dominulus nests can be started by one or more nest founding queens (foundresses). Consequently, there is much interest in the specific benefits that induce cooperation among foundresses. Here, we experimentally demonstrate one major benefit of cooperation, namely that multiple foundresses increase colony productivity. This increase is close to the value predicted by subtracting the productivity of undisturbed single-foundress colonies from the productivity of undisturbed multiple-foundress colonies. However, we found no evidence that an associating foundress' contribution to colony growth is preserved if she disappears (assured fitness returns). Our correlational data suggest that cooperation provides survival benefits, multiple-foundress colonies are more likely to survive to produce offspring than are single-foundress colonies, and individual foundresses in multiple-foundress groups are less likely to disappear before worker emergence than foundresses nesting alone. Therefore, association provides substantial productivity and survival benefits for cooperating foundresses. Copyright 2003.

Paper wasps of the family Vespidae exhibit a wide range of social lifestyles, from facultative eusocial groups to highly ritualistic swarm-founding societies. Even so, adult caste flexibility is widespread throughout the eusocial tribes. Thus, a common endocrine mechanism for caste determination and maintenance in paper wasps is expected, with Polistes dominula serving as a model for the study of mechanisms controlling phenotypic plasticity. In P. dominula, juvenile hormone (JH) and ecdysteroids have been shown to have important caste-determining functions, are important for reproductive growth, and correlate with hydrocarbon signals on the cuticle. Yet research on swarm-founding vespids has shown that JH functions are surprisingly labile, begging the question as to how conserved JH functions are within Polistes, a non-swarming genus. Here, we compared the JH and ecdysteroid titers, cuticular hydrocarbon (CHC) profiles, and prospective visual signals of dominance of free-living foundresses from two sympatric and closely related species of Polistes in South Africa, the indigenous Polistes smithii and the invasive P. dominula. In contrast to P. dominula, neither the JH titer nor the CHC profile was linked to dominance or reproduction in P. smithii, and in both species, hemolymph ecdysteroids were essentially absent. Moreover, many of the relationships between hormones, reproduction, dominance, and social signals in P. dominula are in contrast to studies performed on northern hemisphere populations. The divergence of endocrine and chemical profiles within Polistes offers an unforeseen opportunity to study the evolution of proximate mechanisms underlying phenotypic plasticity.

In temperate climates, female paper wasps typically initiate new colonies in the spring. Several nest-founding tactics have been documented in Polistes species, including solitary nest initiation, joining a cooperative association, usurping an existing nest, or adopting an abandoned nest. Occasionally, exceptionally large groups of females have also been found reusing nests from the previous season. Here we report this phenomenon in introduced populations of the Eurasian species Polistes dominulus. We describe in detail the demographic and genetic characteristics of one such spring colony from Los Angeles, California, USA, which was collected with 84 associated adults and all stages of developing brood in its 613 cells. Genetic and morphological data indicate the presence of multiple reproductively active females of varying relatedness, as well as many nonbreeding females, including probable early-produced offspring. Despite some evidence of chaotic social conditions, the colony appeared to have been highly productive. Additional observations of similar colonies are needed to determine how control is maintained within such a large breeding aggregation.

Polistes versicolor (Olivier) (Hymenoptera: Vespidae) colonies are easily found in anthropic environments; however there is little information available on biological, ecological and behavioral interactions of this species under these environmental conditions. The objective of this work was to characterize the foundation pattern, the productivity, and the success of colonies of P. versicolor in anthropic environments. From August 2003 to December 2004, several colonies were studied in the municipal district of Juiz de Fora, Southeastern Brazil. It was possible to determine that before the beginning of nest construction the foundress accomplishes recognition flights in the selected area, and later begins the construction of the peduncle and the first cell. As soon as new cells are built, the hexagonal outlines appear and the peduncle is reinforced. Foundation of nests on gypsum plaster was significantly larger (p < 0.0001; χ2 test) in relation to the other types of substrate, revealing the synantropism of the species. On average, the P. versicolor nest presents 244.2 ± 89.5 (100–493) cells and a medium production of 171.67 ± 109.94 (37–660) adults. Cells that produced six individuals were verified. Usually, new colonies were founded by an association of females, responsible for the success of 51.5%. Although these results enlarge knowledge on the foundation pattern of P. versicolor in anthropic environments, other aspects of the foundation process require further investigation.

The insectivorous tree swallow was chosen as an indicator species to investigate the uptake of pulp mill-related chlorinated hydrocarbons from emergent aquatic insects. Nest box populations were monitored for reproductive success at locations upstream and downstream of pulp mills on two river systems in British Columbia, Canada. Also, 16-day-old nestlings were collected and analysed for polychlorinated biphenyls (PCBs), polychlorinated dibenzo-p-dioxins (PCDDs) and dibenzofurans (PCDFs), pesticides, chlorophenols and chloroguaiacols. Most reproductive parameters were not different between populations at upstream and downstream locations, and were equivalent to or greater than those recorded for other tree swallow populations. Chlorinated hydrocarbon contamination of nestlings was low at all sites. The highest tissue concentrations were detected downstream of pulp mills on the Fraser River, where PCDD and PCDF patterns along with the presence of pentachlorophenol (PCP) suggested that the primary source of contaminants was past use of PCP for timber preservation. Although the absolute tissue concentration of contaminants was less, the toxic concentration (as estimated with I-TEQs) was greatest in nestlings downstream of a pulp mill on the Thompson River. The proportionately larger contributions from 2,3,7,8-TCDF and PCB-77 elevated these TEQs in comparison to other populations. Nest success was the one reproductive measure that showed substantial reductions in downstream populations on both rivers; however, there was little indication that nest failures were the direct result of contamination. Failures were largely due to parental abandonment, and, while poor parental attentiveness and nest abandonment have been associated with chlorinated hydrocarbon tissue concentrations in other studies, our 1-year assessment was insufficient to establish any link with pulp mill effluent exposure. Nestling growth models showed some subtle differences in growth patterns between nest box populations on the two rivers, but an association with pulp mill effects on aquatic insect prey availability was not established.

The first years of this century have seen significant advances in integrating the many perspectives on what it will take to achieve a healthy and sustainable future. Intense activity among agencies worldwide has produced a stream of reviews of the global condition. These include the combined United States Research Council (1999); a consortium of the United Nations Development Programme, United Nations Environment Programme (UNEP), World Bank, and World Resources Institute (2000); the Millennium Development Goals (2000). United Nations Environment Program (2003); the Global Reporting Initiative (2006); and Millennium Ecosystem Assessment (2005). Each review has deplored a lack of adequate follow-up action on the implications of previous reports. Continuing concern about the state of the planet and the associated risks to health (Millennium Ecosystem Assessment, 2005; Corvalen et al., 2005) has increased the challenge to deliver an effective response. That success is possible has been demonstrated in the Montreal protocols, in which United Nations members continually monitor and control their use of chlorofluorocarbons and so slow the disintegration of the planet s ozone layer (United Nations Environment Programme, 1999). Local government authorities agreement to reduce their carbon emissions in the face of global warming is beginning to show results (ICLEI, 2002). Such effective responses are still far from established practice. There is an inability, first, to agree on, and then to implement, protocols controlling global environmental hazards; for instance, the Kyoto protocols for reducing national carbon dioxide emissions have only just established a framework (United Nations Environment Program, 2006). At the local scale, the management of ecosystem resources tends not to take sufficient account of the needs for long-term sustainability (Corvalen et al., 2005). These precedents confirm that there is much yet to be learned on taking action in response to scientific reports on the state of the world. In this essay, we report on a workshop on that theme held at the EcoHealth ONE meeting, in Madison, Wisconsin in October 2006. The workshop reviewed the three initiatives below with the aim of determining how EcoHealth actions can best be guided in the future. One, a master plan has been provided by the suite of eight United Nations Millennium Development Goals, formed from the synthesis of national commitments made separately at conferences and summits during the 1990s Published online: February 7, 2007 Correspondence to: Valerie A. Brown, e-mail: val.brown@anu.edu.au EcoHealth 4, 95–98, 2007 DOI: 10.1007/s10393-007-0082-8

Social insect colonies often have one or a few queens. How these queens maintain their reproductive monopoly, when other colony members could gain by sharing in the reproduction, is not generally known. DNA microsatellite genotyping is used to determine reproductive interests of various classes of colony members in the paper wasp, Polistes annularis. The relatedness estimates show that the best outcome for most individuals is to be the reproductive egg-layer. For workers, this depends on the sex of offspring: they should prefer to lay their own male eggs, but are indifferent if the queen lays the female eggs. The next-best choice is usually to support the current queen. As a rule, subordinates and workers should prefer the current queen to reproduce over other candidates (though subordinates have no strong preference for the queen over other subordinates, and workers may prefer other workers as a source of male eggs). This result supports the theory that reproductive monopoly stems from the collective preferences of non-reproductives, who suppress each other in favor of the queen. However, we reject the general hypothesis of collective worker control in this species because its predictions about who should succeed after the death of the present queen are not upheld. The first successor is a subordinate foundress even though workers should generally prefer a worker successor. If all foundresses have died, an older worker succeeds as queen, in spite of a collective worker preference for a young worker. The results support the previous suggestion that age serves as a conventional cue serving to reduce conflict over queen succession.

We observed in the field how resident females of the social wasp, Polistes fuscatus, treated female nestmates, non-nestmate first cousins, and unrelated non-nestmates. Behavioral observations were made 1, 3, 4, 6, 7, 8, and 11 weeks after the emergence of a colony's first workers. Resident females were highly tolerant of female nestmates but highly intolerant of unrelated non-nestmates throughout the post-worker-emergence phase of the colony cycle. First cousins were treated either highly tolerantly or highly intolerantly, although most cousins (∼ 63%) were treated highly intolerantly. The treatment of cousins suggests that they are treated either tolerantly (like nestmates) or intolerantly (like unrelated non-nestmates) depending on whether they fall above or below an acceptance threshold, respectively. Resident females did not significantly change their tolerance nestmates, first cousins, and unrelated females between different observation periods. However, all conspecifics were generally treated the most intolerantly 11 weeks post-worker-emergence, a time in the colony cycle when combs no longer contain brood. This decreased tolerance and its associated variance patterns late in the colony cycle are consistent with a more restrictive acceptance threshold. The optimal level of the acceptance threshold appears to be a compromise between the fitness consequences of accepting unrelated non-nestmates and rejecting one's nestmates. In a laboratory study, resident females were equally tolerant of female nestmates on and off the comb but significantly less tolerant of unrelated females on the comb than off the comb. Thus, females can modify their tolerance rapidly in different recognition contexts. In a second laboratory study, resident females were equally intolerant of unrelated females, regardless of whether they were from colonies 50 m or 50 km distant.

We watch two children playing the game ‘‘Rock, Paper, Scissors.’’ This game begins as one of random chance, but it rapidly evolves into a mesmerizing interaction of strategy, wit, and memory. The hand is transformed into a weapon, an attack, a defense. Now cutting, now wrapping, now blunting. Fingers become blades; fists become rock; palms become paper. In this issue’s cover art, one part of a triptych by the renowned Mexican artist Abel Vazquez, a most curious fish brings new meaning to that childhood game. This pajaro is a mythical beast, a transformed animal, a bizarre chimera. A bird with human qualities. A bird with a bowler hat. A bird with the tail of a fish. This striking watercolor is a contemporary take on the alebrije figurines first created by the celebrated Mexican artist Pedro Linares in 1936. It was in this year, so the story goes, that Pedro Linares became ill and, while lying feverish in bed, he dreamt of a surreal forest where animals transformed into chimeric forms, each shouting the word ‘‘alebrijes!’’ When Linares awoke, he picked up the nearest material—paper— and molded figures of these chimeric forms, painting them in garish colors remembered through the distortion of severe illness. Linares’ bestiary is now created by his sons and grandsons and a host of ‘‘cartoneros’’ across Mexico. Beautiful winged fish with legs; dragons grasping at decorated skeletons; bejeweled, deformed toad-headed agamids. Like the mythical Chimera of the 9th century B.C. manuscripts by Homer (Iliad, VI and XVI) and Hesiod (The Theogony, ll. 306–332), these beasts are a thing of fascination, curiosity, and fear for humans. While many, like our cover’s image, are harmless, others represent a magnified ability to destroy and kill. A threat to our very existence. A subliminal stab to our psyche. And so, we return to Mexico, to a more dramatic version of ‘‘Rock, Paper, Scissors’’ played out very recently. This time, it’s a game of genetic reassortment—random acts of chemical bonding. But, as in the children’s game, much more than chance guides the outcome. Given the right conditions—high densities of livestock, international trade and travel, and a diversity of migrating birds—we find ourselves in a similar mesmerizing battle, one that pitches our wits against the evolution of our own chimeric H1N1 creation. What can we learn from Greek mythology that might have bearing on the defeat on this modern, man-made chimera? In mythical times, the Greek gods sent a hero, Bellerophon, to destroy the Chimera. He did this by tipping his spear with lead, melting this in the beast’s fiery breath, and skewering the animal and sealing its guts (Iliad, XVI). A gory tale, perhaps, but also a measure of our own fate? With H1N1 influenza now global, and a simmering caseload in Australia ready to burst forth on the Northern Hemisphere’s winter flu season, we are in unknown territory as we scramble for a vaccine. We analyze our chimera’s weapons, its fiery breath, and we design our spears accordingly. But our shifting foe is not the known entity that the ancient Chimera was. Driven by our own actions, it is able to respond to our every move and transform again— evolving resistance, changing virulence. And so, returning to our Mexican artist Abel Vazquez, we watch as he puts down his brush, and silently we wait. This artist is still painting. This triptych is not complete. EcoHealth 6, 159–160, 2009 DOI: 10.1007/s10393-009-0245-x

Laboratory studies on overwintered foundresses of Polistes gallicus show that differences in the endocrine activity are mainly responsible for achieving the dominance rank. Females that became dominant had either larger corpora allata or more developed ovaries than subordinate females. Body size did not contribute to dominance rank. Since a correlation exists between the length of terminal oocytes and the ecdysteroid titre in haemolymph as well as between the volume of corpora allata and the synthesis of juvenile hormone, dominant behaviour is thought to depend upon an elevated hormone titre in haemolymph. Injections of juvenile hormone (JHI) and 20-hydroxyecdysone, separately and simultaneously, significantly increased the probability that the treated female would be the dominant female of a test pair. After a hierarchy has been established, endocrine activity in subordinate foundresses is inhibited by the dominant foundress that then monopolizes reproduction.

Maternal effects should be especially likely when mothers actively provision offspring with resources that influence offspring phenotype. In cooperatively breeding and eusocial taxa, there is potential for parents to strategically manipulate offspring phenotype in their own interests. Social insect queens are nearly always larger than their worker offspring, and queens could benefit by producing small daughter workers in several ways. If queens use aggression to dominate or coerce workers, a queen producing small workers might minimize potential conflict or competition from her offspring. In addition, because of the trade-off between the number of workers she is able to produce and their individual size, a queen may produce small workers to optimize colony work effort. In this study, we investigate why queens of the primitively eusocial paper wasp Polistes gallicus limit the size of their workers. We created queen–worker size mismatches by cross-fostering queens between nests. We then tested whether the queen–worker size difference affects worker foraging and reproductive effort, or the amount of aggression in the group. Some of our results were consistent with the idea that queens limit worker size strategically: small workers were no less successful foragers, so that producing a larger number of smaller workers may overall increase queen fitness. We found that queens were less likely to attack large workers, perhaps because attempting to coerce large workers is riskier. However, larger workers did not forage less, did not invest more in ovarian development, and were not more aggressive themselves. There was therefore little evidence overall that queens limit conflict by producing smaller workers. Significance statement In social animals, parents might manipulate phenotypic traits of their offspring in their own interests. In paper wasps (Polistes), the first offspring produced are smaller than the queen and become workers: instead of founding their own nests, they stay and help their mother to rear new queens and males. We investigated whether P. gallicus queens could benefit by producing small daughter workers by using cross-fostering to create size mismatches between queens and their offspring. We then recorded foraging activity, reproductive effort, and aggression on nests. Queens were less likely to attack larger workers, but overall, there was limited evidence of size-based queen–worker conflict. However, because small workers were no less successful foragers, producing a larger number of smaller workers may optimize colony work effort.

Abstract The Duke of Burgundy butterfly (Hamearis lucina) is known to have specific habitat requirements for its larval foodplants. However, no studies have yet investigated whether these preferences vary over time or in relation to climate, and there is a paucity of data on whether management on reserves can replicate preferred conditions. Here, we build upon existing research to confirm which characteristics Duke of Burgundy prefer for their larval foodplants, whether preferences remain consistent across years, and whether conservation management on reserves can replicate these conditions. Fieldwork was carried out at Totternhoe Quarry Reserve, a chalk grassland site in Bedfordshire, UK. Confirming previous research, we found that large Primula plants in dense patches were chosen for oviposition, but that once chosen there was no preference to lay eggs on a plant’s largest leaf. Chosen foodplants were also more sheltered and in closer proximity to scrub than their controls. However, at a finer scale, we found little evidence for any preference based on differences in microclimate, or vegetation height immediately surrounding the plants. This suggests features that alter microclimatic conditions at a larger scale are relatively more important for determining the suitability of oviposition sites. Nearly all preferences remained consistent over time and did not vary between years. Management of scrub on the reserve was able to reproduce some preferred habitat features (high plant density), but not others (large plant size). Implications for insect conservation The consistency of findings across years, despite inter-annual variation in temperature, rainfall and number of adults, indicates that the Duke of Burgundy is conservative in its foodplant choice, highlighting its need for specific habitat management. Targeted management for foodplants could form part of a tractable set of tools to support Duke of Burgundy numbers on reserves, but a careful balance is needed to avoid scrub clearance leaving plants in sub-optimal conditions.

Cheliceral chelal design in free-living astigmatid mites (Arthropoda: Acari) is reviewed within a mechanical model. Trophic access (body size and cheliceral reach) and food morsel handling (chelal gape and estimated static adductive crushing force) are morphologically investigated. Forty-seven commonly occurring astigmatid mite species from 20 genera (covering the Acaridae, Aeroglyphidae, Carpoglyphidae, Chortoglyphidae, Glycyphagidae, Lardoglyphidae, Pyroglyphidae, Suidasiidae, and Winterschmidtiidae) are categorised into functional groups using heuristics. Conclusions are confirmed with statistical tests and multivariate morphometrics. Despite these saprophagous acarines in general being simple ‘shrunken/swollen’ versions of each other, clear statistical correlations in the specifics of their mechanical design (cheliceral and chelal scale and general shape) with the type of habitat and food consumed (their ‘biome’) are found. Using multivariate analyses, macro- and microsaprophagous subtypes are delineated. Relative ratios of sizes on their own are not highly informative of adaptive syndromes. Sympatric resource competition is examined. Evidence for a maximum doubling of approximate body volume within nominal taxa is detected but larger mites are not more ‘generalist’ feeding types. Two contrasting types of basic ‘Bauplan’ are found differing in general scale: (i) a large, chunk-crunching, ‘demolition’-feeding omnivore design (comprising 10 macrosaprophagous astigmatid species), and (ii) a small selective picking, squashing/slicing or fragmentary/‘plankton’ feeding design (which may indicate obligate fungivory/microbivory) comprising 20 microsaprophagous acarid-shaped species. Seventeen other species appear to be specialists. Eleven of these are either: small (interstitial/burrowing) omnivores—or a derived form designed for processing large hard food morsels (debris durophagy, typified by the pyroglyphid Dermatophagoides farinae), or a specialist sub-type of particular surface gleaning/scraping fragmentary feeding. Six possible other minor specialist gleaning/scraping fragmentary feeders types each comprising one to two species are described. Details of these astigmatid trophic-processing functional groups need field validation and more corroborative comparative enzymology. Chelal velocity ratio in itself is not highly predictive of habitat but with cheliceral aspect ratio (or chelal adductive force) is indicative of life-style. Herbivores and pest species are typified by a predicted large chelal adductive force. Pest species may be ‘shredders’ derived from protein-seeking necrophages. Carpoglyphus lactis typifies a mite with tweezer-like chelae of very feeble adductive force. It is suggested that possible zoophagy (hypocarnivory) is associated with low chelal adductive force together with a small or large gape depending upon the size of the nematode being consumed. Kuzinia laevis typifies an oophagous durophage. Functional form is correlated with taxonomic position within the Astigmata—pyroglyphids and glycyphagids being distinct from acarids. A synthesis with mesostigmatid and oribatid feeding types is offered together with clarification of terminologies. The chelal lyrifissure in the daintiest chelicerae of these astigmatids is located similar to where the action of the chelal moveable digit folds the cheliceral shaft in uropodoids, suggesting mechanical similarities of function. Acarid astigmatids are trophically structured like microphytophagous/fragmentary feeding oribatids. Some larger astigmatids (Aleuroglyphus ovatus, Kuzinia laevis, Tyroborus lini) approximate, and Neosuidasia sp. matches, the design of macrophytophagous oribatids. Most astigmatid species reviewed appear to be positioned with other oribatid secondary decomposers. Only Dermatophagoides microceras might be a primary decomposer approximating a lichenivorous oribatid (Austrachipteria sp.) in trophic form. Astigmatid differences are consilient with the morphological trend from micro- to macrophytophagy in oribatids. The key competency in these actinotrichid mites is a type of ‘gnathosomisation’ through increased chelal and cheliceral height (i.e., a shape change that adjusts the chelal input effort arm and input adductive force) unrestricted by the dorsal constraint of a mesostigmatid-like gnathotectum. A predictive nomogram for ecologists to use on field samples is included. Future work is proposed in detail.

Commercially-reared bumblebee colonies provide pollination services to numerous crop species globally. These colonies may harbour parasites which can spill-over to wild bee species. However, the potential for parasites to spread from wild to commercial bumblebees, which could then lead to parasite spill-back, is poorly understood. To investigate this, parasite-free commercial Bombus terrestris audax colonies, which are used commercially for strawberry pollination, were placed into seasonal strawberry crops for either 6- or 8-week blocks across two key time periods, early spring and early summer. Bumblebees were removed from colonies weekly and screened for the presence of parasites. In the early spring placement, only one parasite, the highly virulent neogregarine Apicystis bombi, was detected at a low prevalence (0.46% across all bees screened). In contrast, all colonies placed in the crop in the early summer became infected. A trypanosome, Crithidia bombi, and A. bombi were the most prevalent parasites across all samples, reaching peak prevalence in screened bees of 39.39% and 18.18% respectively at the end of the experimental period. The prevalence of A. bombi was greater than most UK records from wild bumblebees, suggesting that commercial colonies could enhance levels of A. bombi infection in wild bees through spill-back. Studies on larger geographical scales with different commercial colony densities are required to fully assess spill-back risk. However, seasonal management, to minimise spill-back opportunities, and treatment of commercial colonies to prevent infection, could be implemented to manage the potential risks of parasite spill-back to wild bees.Implications for insect conservation Our results show that commercial bumblebee populations do pick up infections, most likely from wild bees, and that these infections can reach prevalences where they may pose a threat to wild bees via parasite spill-back. More research is required to clarify the extent of this potential threat.

Major histocompatibility complex (MHC) genes have been widely studied to assess the immunological fitness and evolutionary adaptation of animal populations. Among the Canidae, the raccoon dog’s adventurous nature, omnivorous behavior, and high variability of intracellular pathogens make it ideal to study selection on MHC class I in a non-model canid species. Here, we examined allelic diversity and evolutionary patterns of MHC class I genes in the raccoon dog (Nyctereutes procyonoides). We identified 48 novel MHC class I alleles from 31 raccoon dogs from Japan and Russia. Some alleles were geographically restricted, whereas others were widely distributed across the species’ range. The rate of non-synonymous substitutions was greater than that of synonymous substitutions for both exon 2 and exon 3 encoding α1 and α2 domains, respectively, in the α chain of the MHC class I protein. Positively selected sites at the amino acid level were evident in both the α1 and α2 domains, and a recombination breakpoint was found in exon 3. Bayesian phylogenetic trees showed no evidence of trans-species polymorphism (TSP) with alleles from carnivoran species in other families but did detect TSP between raccoon dogs and the domestic dog, Canis familiaris, indicative of long-term balancing selection in canids. Our results indicate that the extensive allelic diversity of MHC class I in Japanese and Russian raccoon dogs has been influenced and maintained by pathogen-driven positive selection, recombination, and long-term balancing selection. Supplementary Information The online version contains supplementary material available at 10.1007/s13364-021-00561-y.

The founders of a social group or colony have the potential to greatly influence the success or failure of the societies that they initiate. Whether through their genetic contribution, or through behavioral maternal effects related to temperament, resource acquisition, or brood care, the traits of founders deserve special consideration. The queens of many social insects are ideally suited to address questions relating to the importance of founder traits due to her solitary involvement in performing a multitude of tasks necessary to produce a functioning colony. While it has been suggested that a queen’s behavioral phenotype might contribute to differences in colony success, no study has yet demonstrated these links in situ. Here, we use the singly founding (haplometrotic) paper wasp Polistes metricus to examine whether queen personality, measured shortly after colony founding (pre-emergent phase), and morphology, can predict colony size (a proxy for queen fitness) and parasite load in the wild. We found that larger and bolder queens gave rise to larger colonies than smaller and shyer queens, and there was no relationship between queen personality and parasite load. Differences in queen traits therefore appear to be a major determinant of colony success under field conditions. In social species, the personality composition of groups has been shown to be a large determinant of group behavior, which in turn can determine group performance and survival. In this study, we explore the degree to which the behavioral tendencies of key group members—wasp queens—influence the performance of their descendant societies. We tested the behavioral tendencies and morphology of Polistes metricus paper wasp queens in the early spring, placed them back in the field in nest boxes, and recorded each nest’s cell count at the end of the season—a reliable proxy for queen fitness. We found that larger and bolder queens gave rise to larger descendant colonies on average than their smaller, shyer counterparts. Individual variation in queen traits therefore appears to be a determinant of colony success under field conditions.

A model based upon mechanics is used in a re-analysis of historical acarine morphological work augmented by an extra seven zoophagous mesostigmatid species. This review shows that predatory mesostigmatids do have cheliceral designs with clear rational purposes. Almost invariably within an overall body size class, the switch in predatory style from a worm-like prey feeding (‘crushing/mashing’ kill) functional group to a micro-arthropod feeding (‘active prey cutting/slicing/slashing' kill) functional group is matched by: an increased cheliceral reach, a bigger chelal gape, a larger morphologically estimated chelal crunch force, and a drop in the adductive lever arm velocity ratio of the chela. Small size matters. Several uropodines (Eviphis ostrinus, the omnivore Trachytes aegrota, Urodiaspis tecta and, Uropoda orbicularis) have more elongate chelicerae (greater reach) than their chelal gape would suggest, even allowing for allometry across mesostigmatids. They may be: plesiosaur-like high-speed strikers of prey, scavenging carrion feeders (like long-necked vultures), probing/burrowing crevice feeders of cryptic nematodes, or small morsel/fragmentary food feeders. Some uropodoids have chelicerae and chelae which probably work like a construction-site mechanical excavator-digger with its small bucket. Possible hoeing/bulldozing, spore-cracking and tiny sabre-tooth cat-like striking actions are discussed for others. Subtle changes lead small mesostigmatids to be predator–scavengers (mesocarnivores) or to be predator–fungivores (hypocarnivores). Some uropodines (e.g., the worm-like prey feeder Alliphis siculus and, Uropoda orbicularis) show chelae similar in design to astigmatids and cryptostigmatids indicating possible facultative saprophagy. Scale matters—obligate predatory designs (hypercarnivory) start for mesostigmatids with chelal gape > 150 μm and cheliceral reach > 350 μm (i.e., about 500–650 μm in body size). Commonality of trophic design in these larger species with solifugids is indicated. Veigaia species with low chelal velocity ratio and other morphological strengthening specialisms, appear specially adapted in a concerted way for predating active soft and fast moving springtails (Collembola). Veigaia cerva shows a markedly bigger chelal gape than its cheliceral reach would proportionately infer suggesting it is a crocodile-like sit-and-wait or ambush predator par excellence. A small chelal gape, low cheliceral reach, moderate velocity ratio variant of the worm-like feeding habit design is supported for phytoseiid pollenophagy. Evidence for a resource partitioning model in the evolution of gnathosomal development is found. A comparison to crustacean claws and vertebrate mandibles is made. Alliphis siculus and Rhodacarus strenzkei are surprisingly powerful mega-cephalics for their small size. Parasitids show a canid-like trophic design. The chelicera of the nematophagous Alliphis halleri shows felid-like features. Glyphtholaspis confusa has hyaena-like cheliceral dentition. The latter species has a markedly smaller chelal gape than its cheliceral reach would suggest proportionately, which together with a high chelal velocity ratio and a high estimated chelal crunch force matches a power specialism of feeding on immobile tough fly eggs/pupae by crushing (durophagy). A consideration of gnathosomal orientation is made. Predatory specialisms appear to often match genera especially in larger mesostigmatids, which may scale quite differently. Comparison to holothyrids and opilioacarids indicates that the cheliceral chelae of the former are cutting-style and those of the latter are crushing-style. A simple validated easy-to-use ‘2:1 on’ predictive algorithm of feeding habit type is included based on a strength-speed tradeoff in chelal velocity ratio for ecologists to test in the field.

Niche construction refers to the modification of selective environments by organisms. Theoretical and empirical studies of niche construction are increasing in importance as foci in evolutionary ecology. This special edition presents theoretical and empirical research that illustrates the significance of niche construction to the field. Here we set the scene for the following papers by (1) discussing the history of niche construction research, (2) providing clear definitions that distinguish niche construction from related concepts such as ecosystem engineering and the extended phenotype, (3) providing a brief summary of the findings of niche construction research, (4) discussing the contribution of niche construction and ecological inheritance to (a) expanded notions of inheritance, and (b) the extended evolutionary synthesis, and (5) briefly touching on some of the issues that underlie the controversies over niche construction. Publisher PDF