Intercontinental chemical variation in the invasive ant Wasmannia auropunctata (Roger) (Hymenoptera Formicidae): a key to the invasive success of a tramp species

Unicoloniality emerges as a feature that characterizes successful invasive species. Its underlying mechanism is reduced intraspecific aggression while keeping interspecific competitiveness. To that effect, we present here a comparative behavioural and chemical study of the invasive ant Wasmannia auropunctata in parts of its native and introduced ranges. We tested the hypothesis that introduced populations (New Caledonia archipelago) have reduced intraspecific aggression relative to native populations (e.g., Ilhéus area, Brazil) and that this correlates with reduced variability in cuticular hydrocarbons (CHCs). As predicted, there was high intraspecific aggression in the Brazilian populations, but no intraspecific aggression among the New Caledonian populations. However, New Caledonian worker W. auropunctata remained highly aggressive towards ants of other invasive species. The chemical data corresponded with the behaviour. While CHCs of ants from the regions of Brazil diverged, the profiles of ants from various localities in New Caledonia showed high uniformity. We suggest that in New Caledonia W. auropunctata appears to behave as a single supercolony, whereas in its native range it acts as a multicolonial species. The uniformity of recognition cues in the New Caledonia ants may reflect a process whereby recognition alleles became fixed in the population, but may also be the consequence of a single introduction event and subsequent aggressive invasion of the ecosystem. Chemical uniformity coupled with low intraspecific but high interspecific aggression, lend credence to the latter hypothesis.     

Managing dependencies in forest offset projects: toward a more complete evaluation of reversal risk

Although forest carbon offsets can play an important role in the implementation of comprehensive climate policy, they also face an inherent risk of reversal. If such risks are positively correlated across projects, it can affect the integrity of larger project portfolios and potentially the entire offsets program. Here, we discuss three types of risks that could affect forest offsets—fat tails, micro-correlation, and tail dependence—and provide examples of how they could present themselves in a forest offset context. Given these potential dependencies, we suggest several new risk management approaches that take into account dependencies in reversal risk across projects and which could help guard the climate integrity of an offsets program. We also argue that data collection be included as an integral part of any offsets program so that disturbance-related dependencies may be identified and managed as early and to the greatest extent possible.     

Allometric shape change of the lower pharyngeal jaw correlates with a dietary shift to piscivory in a cichlid fish

The morphological versatility of the pharyngeal jaw of cichlid fishes is assumed to represent a key factor facilitating their unparalleled trophic diversification and explosive radiation. It is generally believed that the functional design of an organism relates to its ecology, and thus, specializations to different diets are typically associated with distinct morphological designs, especially manifested in the cichlids’ pharyngeal jaw apparatus. Thereby, the lower pharyngeal jaw (LPJ) incorporates some of the most predictive features for distinct diet-related morphotypes. Thus, considering that piscivorous cichlids experience an ontogenetic dietary shift from typically various kinds of invertebrates to fish, concomitant morphological changes in the LPJ are expected. Using Lepidiolamprologus elongatus, a top predator in the shallow rocky habitat of Lake Tanganyika, as model, and applying geometric and traditional morphometric techniques, we demonstrate an allometric change in ontogenetic LPJ shape development coinciding with the completion of the dietary shift toward piscivory. The piscivorous LPJ morphotype is initiated in juvenile fish by increasing elongation and narrowing of the LPJ and—when the fish reach a size of 80–90 mm standard length—further refined by the elongation of the posterior muscular processes, which serve as insertion for the fourth musculus levator externus. The enlarged muscular processes of the fully mature piscivorous morphotype provide for the construction of a powerful lever system, which allows the large individuals to process large prey fish and rely on exclusive piscivory.     

The attractiveness fragment—AFLP analysis of local adaptation and sexual selection in a caeliferan grasshopper, <Emphasis Type="Italic">Chorthippus biguttulus</Emphasis>

Genetic variability among males is a necessary precondition for the evolution of female choice based on indirect genetic benefits. In addition to mutations and host–parasite cycles, migration of locally adapted individuals offers an explanation for the maintenance of genetic variability. In a previous study, conducting a reciprocal transplant experiment on a grasshopper, Chorthippus biguttulus, we found that environmental conditions significantly influenced not only body condition but also an important trait of male calling song, the amplitude of song. Although not significant, all other analysed physical and courtship song traits and attractiveness were superior in native than in transferred males. Thus, we concluded that local adaptation has a slight but consistent influence on a range of traits in our study populations, including male acoustic attractiveness. In our present study, we scanned male grasshoppers from the same two populations for amplification fragment length polymorphism (AFLP) loci connected with acoustic attractiveness to conspecific females. We found greater differences in allele frequencies between the two populations, for some loci, than are expected from a balance between drift and gene flow. These loci are potentially connected with locally adapted traits. We examined whether these alleles show the proposed genotype environment interaction by having different associations with attractiveness in the two populations. One locus was significantly related to sexual attractiveness; however, this was independent of the males’ population affiliation. Future research on the evolution of female choice will benefit from knowledge of the underlying genetic architecture of male traits under intraspecific sexual selection, and the ‘population genomics’ approach can be a powerful tool for revealing this structure.     

Run for your life,but bite for your rights? How interactions between natural and sexual selection shape functional morphology across habitats

A central issue in evolutionary biology is how morphology, performance, and habitat use coevolve. If morphological variation is tightly associated with habitat use, then differences in morphology should affect fitness through their effect on performance within specific habitats. In this study, we investigate how evolutionary forces mold morphological traits and performance differently given the surrounding environment, at the intraspecific level. For this purpose, we selected populations of the lizard Podarcis bocagei from two different habitat types, agricultural walls and dunes, which we expected to reflect saxicolous vs ground-dwelling habits. In the laboratory, we recorded morphological traits as well as performance traits by measuring sprint speed, climbing capacity, maneuverability, and bite force. Our results revealed fast-evolving ecomorphological variation among populations of P. bocagei, where a direct association existed between head morphology and bite performance. However, we could not establish links between limb morphology and locomotor performance at the individual level. Lizards from walls were better climbers than those from dunes, suggesting a very fast evolutionary response. Interestingly, a significant interaction between habitat and sex was detected in climbing performance. In addition, lizards from dunes bit harder than those from walls, although sexual differentiation was definitely the main factor driving variation in head functional morphology. Taking into account all the results, we found a complex interaction between natural and sexual selection on whole-organism performance, which are, in some cases, reflected in morphological variation.     

Estimating the magnitude of morphoscapes: how to measure the morphological component of biodiversity in relation to habitats using geometric morphometrics

Ecological indicators are currently developed to account for the different facets of loss of biological diversity due to direct or indirect effects of human activities. Most ecological indicators include species richness as a metric. Others, such as functional traits and phylogenetic diversity, account for differences in species, even when species richness is the same. Here, we describe and apply a different indicator, called morphoscape dimension, accounting for morphological variability across habitats in a geographical region. We use the case of ground beetles (Coleoptera: Carabidae) in four different habitats in the Po Plain in Northern Italy to exemplify how to quantify the magnitude of the morphological space (i.e. the dimension of the morphoscape) occupied by the species in each habitat using geometric morphometrics. To this aim, we employed a variety of metrics of morphological disparity related to univariate size, and more complex multivariate shape and form. Our ‘proof of concept’ suggests that metrics assessing size and form might largely tend to simply mirror the information provided by species richness, whereas shape morphoscape disparity may be able to account for non-trivial differences in species traits amongst habitats. This is indicated by the woodland morphoscape being on average bigger than that of crops, the most species-rich habitat, despite having almost 20% less species. We conclude suggesting that the analysis of morphoscape dimension has the potential to become a new additional and complimentary tool in the hands of conservation biologists and ecologists to explore and quantify habitat complexity and inform decisions on management and conservation based on a wide set of ecological indicators.     

Consequences of correlations between habitat modifications and negative impact of climate change for regional species survival

While several empirical and theoretical studies have clearly shown the negative effects of climate or landscape changes on population and species survival only few of them addressed combined and correlated consequences of these key environmental drivers. This also includes positive landscape changes such as active habitat management and restoration to buffer the negative effects of deteriorating climatic conditions. In this study, we apply a conceptual spatial modelling approach based on functional types to explore the effects of both positive and negative correlations between changes in habitat and climate conditions on the survival of spatially structured populations. We test the effect of different climate and landscape change scenarios on four different functional types that represent a broad spectrum of species characterised by their landscape level carrying capacity, the local population turnover rates at the patch level (K-strategies vs. r-strategies) and dispersal characterstics. As expected, simulation results show that correlated landscape and climatic changes can accelerate (in case of habitat loss or degradation) or slow down (in case of habitat gain or improvement) regional species extinction. However, the strength of the combined changes depends on local turnover at the patch level, the overall landscape capacity of the species, and its specific dispersal characteristics. Under all scenarios of correlated changes in habitat and climate conditions we found the highest sensitivity for functional types representing species with a low landscape capacity but a high population growth rate and a strong density regulation causing a high turnover at the local patch level.The relative importance of habitat loss or habitat degradation, in combination with climate deterioration, differed among the functional types. However, an increase in regional capacity revealed a similar response pattern: For all types, habitat improvement led to higher survival times than habitat gain, i.e. the establishment of new habitat patches. This suggests that improving local habitat quality at a regional scale is a more promising conservation strategy under climate change than implementing new habitat patches. This conceptual modelling study provides a general framework to better understand and support the management of populations prone to complex environmental changes.     

The European land leech: biology and DNA-based taxonomy of a rare species that is threatened by climate warming

The European land leech Xerobdella lecomtei was discovered in 1868 and is one of the rarest animals on Earth. During the 1960s, several individuals of these approx. 40 mm long, cold-adapted terrestrial annelids that inhabit the moist soils of birch forests around Graz, Austria, were investigated. Only one original research paper has been published on the biology of this species. Between 2001 and 2005, we re-investigated the morphology of preserved specimens and searched for living individuals in their natural habitat that appeared to be intact. We found only one juvenile individual (length approx. 10 mm), indicating that this local leech population became largely extinct over the past four decades. The feeding behaviour of our ‘lonesome George of the annelids’ was studied and is described here in detail. After its death, the Xerobdella individual was used for chemical extraction and molecular studies (deoxyribonucleic acid [DNA] barcoding, based on one gene, the mitochondrial cytochrome c oxidase subunit I). In addition, novel DNA barcodes for a land leech from Madagascar and a recently discovered species from Europe were obtained. Our phylogenetic tree shows that X. lecomtei is not a member of the tropical land leeches (family Haemadipsidae), as previously thought, but represents a separate line of descent (family Xerobdellidae). The decline of the local leech population around Graz correlates with a rise in average summer temperatures of +3°C between 1961 and 2004. This warming led to a drastic reduction in the moisture content of the soil where X. lecomtei lives. We suggest that human-induced climate change without apparent habitat destruction can lead to the extinction of populations of cold-adapted species that have a low colonization ability.     

The role of spatial processes and environmental determinants in microgeographic shell variation of the freshwater snail <Emphasis Type="Italic">Chilina dombeyana</Emphasis> (Bruguière, 1789)

Wildlife data often show spatial organization, demonstrating positive correlations either as a result of processes occurring over the landscape or due to the influence of spatially structured environmental variables. It is, thus, essential to consider non-random spatial structure when evaluating the underlying causes of biological variation. In this study, we analyzed the population structure of Chilina dombeyana shell morphology of 14 populations that are close geographically and belong to the same hydrographic basin. We utilized a variation partitioning approach to evaluate the importance of spatial processes, such as migration, acting over the landscape, and environmental characteristics, including habitat and hydrologic characteristics, and the occurrence of aquatic predators in promoting between population variation. Our results demonstrate spatially structured variation in C. dombeyana shell morphology, with populations living near each other having more similar shell sizes than populations living farther apart. The shell size variation partition indicated that both spatially structured environmental factors and genetic relationships resulting from migration or shared common ancestry may explain this pattern. Shell shape variation, in contrast, was found to be essentially under the influence of non-spatially structured environmental factors, with habitat and water characteristics accounting for about half of the total variation among populations. The large proportion of the variation in shell size that is spatially structured demonstrates that spatial structure on morphological traits might be strong and highlights the need to consider such phenomenon in intraspecific studies of phenotypic evolution.     

Managing climate change risks in New York City’s water system: assessment and adaptation planning

Managing risk by adapting long-lived infrastructure to the effects of climate change must become a regular part of planning for water supply, sewer, wastewater treatment, and other urban infrastructure during this century. The New York City Department of Environmental Protection (NYCDEP), the agency responsible for managing New York City’s (NYC) water supply, sewer, and wastewater treatment systems, has developed a climate risk management framework through its Climate Change Task Force, a government-university collaborative effort. Its purpose is to ensure that NYCDEP’s strategic and capital planning take into account the potential risks of climate change—sea-level rise, higher temperature, increases in extreme events, changes in drought and flood frequency and intensity, and changing precipitation patterns—on NYC’s water systems. This approach will enable NYCDEP and other agencies to incorporate adaptations to the risks of climate change into their management, investment, and policy decisions over the long term as a regular part of their planning activities. The framework includes a 9-step Adaptation Assessment procedure. Potential climate change adaptations are divided into management, infrastructure, and policy categories, and are assessed by their relevance in terms of climate change time-frame (immediate, medium, and long term), the capital cycle, costs, and other risks. The approach focuses on the water supply, sewer, and wastewater treatment systems of NYC, but has wide application for other urban areas, especially those in coastal locations.     

Scale-dependent analysis of an otter–crustacean system in Argentinean Patagonia

The Southern river otter or ‘huillin’, Lontra provocax, is an endangered species endemic of the Andean Patagonian region of Argentina and Chile. It feeds almost exclusively on the genera of macro-crustacea: Aegla and Sammastacus. The aim of this study was to analyse the role of food availability on the huillin’s distribution using a scale-dependent analysis of crustacean and otter distributions. We compared the distributions of otters and macro-crustaceans along a north–south regional gradient, between river basins of northern Patagonia, in an altitudinal gradient within a river basin, and between habitat types within a lake. We investigated the distribution of otters by sign surveys along lake shores, river banks and marine coasts, and of crustaceans using surveys in the water, undigested remains in mink (Mustela vison) scats, presence of external skeletons at the waterside and through interviews with local people. Our results show that there were heterogeneities in the distributions of macro-crustaceans at four scales and these were generally reflected in the distributions of freshwater otters. We conclude that the main factor limiting the distributions of L. provocax in freshwater environments is the availability of macro-crustaceans. This paper shows how scale-dependent type analyses of population distribution serves as a method for identifying key environmental factors for species for which the use of long-term demographies is unfeasible.     

Diverse migration strategies in hoopoes (<Emphasis Type="Italic">Upupa epops</Emphasis>) lead to weak spatial but strong temporal connectivity

The annual cycle of migrating birds is shaped by their seasonal movements between breeding and non-breeding sites. Studying how migratory populations are linked throughout the annual cycle—migratory connectivity, is crucial to understanding the population dynamics of migrating bird species. This requires the consideration not only of spatial scales as has been the main focus to date but also of temporal scales: only when both aspects are taken into account, the degree of migratory connectivity can be properly defined. We investigated the migration behaviour of hoopoes (Upupa epops) from four breeding populations across Europe and characterised migration routes to and from the breeding grounds, location of non-breeding sites and the timing of key migration events. Migration behaviour was found to vary both within and amongst populations, and even though the spatial migratory connectivity amongst the populations was weak, temporal connectivity was strong with differences in timing amongst populations, but consistent timing within populations. The combination of diverse migration routes within populations and co-occurrence on the non-breeding grounds between populations might promote exchange between breeding populations. As a result, it might make hoopoes and other migrating bird species with similar strategies more resilient to future habitat or climatic changes and stabilise population trends.     

Role of winter temperature and climate change on the survival and future range expansion of the hemlock woolly adelgid (Adelges tsugae) in eastern North America

Global climate change has already affected the abundances, range limits, and interactions of many species. The hemlock woolly adelgid (Adelges tsugae), an invasive insect introduced to eastern North America from Japan, has decimated stands of eastern hemlock (Tsuga canadensis) and Carolina hemlock (T. caroliniana) from Georgia to Connecticut. However, its spread across central and northern New England has been slowed substantially by its inability to tolerate cold winter temperatures. Using data from previous lab and field studies collected over the past 17 years, including adelgid spread and overwintering mortality, we first characterize the temperature conditions that may limit adelgid spread. We then show how, in the future, rising winter temperatures due to climate change are likely to remove the conditions currently limiting adelgid spread, and facilitate the northward expansion as more suitable habitat becomes available.     

The evolution of bird migration—a synthesis

We approach the problem of the evolution of bird migration by asking whether migration evolves towards new breeding areas or towards survival areas in the non-breeding season. Thus, we avoid the ambiguity of the usually discussed “southern-home-theory” or “northern-home-theory”. We argue that migration evolved in birds that spread to seasonal habitats through gradual dispersal to enhance survival during the non-breeding season; this in contrast to the alternative idea suggesting that migration evolved towards new breeding areas to increase reproductive success. Our synthesis is based on the threshold model explaining how migratory traits can change rapidly through microevolutionary processes. Our model brings former theories together and explains how bird migration, with the appropriate direction and time program, evolves through selection after genetically non-directed events such as dispersal and colonization. The model does not need the former untested assumptions such as competition as a reason for migration and for the disappearance of sedentary populations or higher reproductive success in temperate breeding areas. Our theory offers answers to questions such as how birds with a southern origin may gradually reach northern latitudes, why migration routes may follow historical expansion routes and why birds leave an area for the non-breeding season and move back instead of breeding on their wintering grounds. The theory proposes gradual change through selection and not sudden changes such as long distance dispersal or mutations and can be applied to migration at all latitudes and in all directions. The scenario provides a reasonable concept to understand most of the existing migratory phenomena on the basis of the ecology and genetics of migratory behaviour.     

Global Carbon Dioxide Emissions Scenarios: Sensitivity to Social and Technological Factors in Three Regions

Carbon dioxide emissions from 1990 to 2100 AD are decomposed into the product of four factors: population size, affluence (measured here as GDP per capita), energy intensity (energy use per unit GDP) and carbon intensity (carbon dioxide emissions per unit energy). These emissions factors are further subdivided into three regions: more developed countries (MDCs), China, and the remaining less developed countries (LDCs). Departures from a baseline scenario (based on IPCC, 1992a — the so-called ‘business-as-usual’ scenario) are calculated for a variety of alternative assumptions concerning the four emissions factors in the three regions. Although the IPCC scenario is called a ‘non-intervention’ scenario, it is shown, for example, that large decreases in energy intensity in China or carbon intensity in MDCs are built into the ‘business as usual’ case — and such large changes vary considerably from region to region. We show what CO₂ emissions would look like if each of these four emissions factors projected in the baseline case somehow remained constant at 1990 levels. Certain factors like energy intensity improvements and long-term population growth in LDCs, or GDP growth and carbon intensity improvements in MDCs, are shown to have a big contribution to cumulative global emissions to 2100 AD, and consequently, changes in these projected factors will lead to significant deviations from baseline emissions. None of the scenarios examined in this analysis seems to indicate that any one global factor is clearly dominant, but cultural, economic, and political costs or opportunities of altering each factor may differ greatly from country to country.     

Explanative power of variables used in species distribution modelling: an issue of general model transferability or niche shift in the invasive Greenhouse frog (Eleutherodactylus planirostris)

The use of species distribution models (SDMs) to predict potential distributions of species is steadily increasing. A necessary assumption when projecting models throughout space or time is that climatic niches are conservative, but recent findings of niche shifts during biological invasion of particular plant and animal species have indicated that this assumption is not categorically valid. One reason for observed shifts may relate to variable selection for modelling. In this study, we assess differences in climatic niches in the native and invasive ranges of the Greenhouse frog (Eleutherodactylus planirostris). We analyze which variables are more ‘conserved’ in comparison to more ‘relaxed’ variables (i.e. subject to niche shift) and how they influence transferability of SDMs developed with Maxent on the basis of ten bioclimatic layers best describing the climatic requirements of the target species. We focus on degrees of niche similarity and conservatism using Schoener's index and Hellinger distance. Significance of results are tested with null models. Results indicate that the degrees of niche similarity and conservatism vary greatly among the predictive variables. Some shifts can be attributed to active habitat selection, whereas others apparently reflect variation in the availability of climate conditions or biotic interactions between the frogs' native and invasive ranges. Patterns suggesting active habitat selection also vary among variables. Our findings evoke considerable implications on the transferability of SDMs over space and time, which is strongly affected by the choice and number of predictors. The incorporation of ‘relaxed’ predictors not or only indirectly correlated with biologically meaningful predictors may lead to erroneous predictions when projecting SDMs. We recommend thorough assessments of invasive species' ecology for the identification biologically meaningful predictors facilitating transferability.     

Collaboration mobilises institutions with scale-dependent comparative advantage in landscape-scale biodiversity conservation

Landscape-scale approaches are emerging as central to ecosystem management and biodiversity conservation globally, triggering the requirement for collaboration between multiple actors and associated risks including knowledge asymmetries; institutional fragmentation; uncertainty; power imbalances; “invisible” slow-changing variables; and entrenched socio-economic inequities. While social science has elucidated some dimensions required for effective collaboration, little is known about how collaboration manages these risks, or of its effects on associated social-ecological linkages. Our analysis of four different Australian contexts of collaboration shows they mobilised institutions matched to addressing environmental threats, at diverse scales across regulatory and non-regulatory domains. The institutions mobilised included national regulatory controls on development that threatened habitat, incentives to farmers for practice-change, and mechanisms that increased resources for on-ground fire and pest management. Knowledge-sharing underpinned effective risk management and was facilitated through the use of boundary objects, enhanced multi-stakeholder peer review processes, interactive spatial platforms, and Aboriginal-driven planning. Institutions mobilised in these collaborations show scale-dependent comparative advantage for addressing environmental threats. The findings confirm the need to shift scientific attention away from theorising about the ideal-scale for governance. We argue instead for a focus on understanding how knowledge-sharing activities across multiple scales can more effectively connect environmental threats with the most capable institution to address these threats.     

Does bird species diversity vary among forest types? A local-scale test in Southern Chile

Birds are the most diverse vertebrate group in Chile, characterized by low species turnover at the country-size scale (high alpha but low beta diversities), resembling an island biota. We tested whether this low differentiation is valid at a local scale, among six forest habitat types. We detected 25 bird species; avifauna composition was significantly different among habitat types, with five species accounting for 60 % of the dissimilarity. We found a higher level of bird assemblage differentiation across habitats at the local scale than has been found at the country-size scale. Such differentiation might be attributed to structural differences among habitats.     

湘江流域传统村落景观基因变异及其分异规律

传统村落在历史、文化、经济、旅游和科学等多个方面价值颇高,保护传统村落的空间格局和文化内涵极为重要。以中国湘江流域上中下游六个典型传统村落为例,以景观基因理论为基础,运用景观基因分析法识别了湘江流域传统村落景观基因,分析了上中下游传统村落显性物质景观基因与隐性非物质文化基因的变异特征,总结了湘江流域传统村落景观基因变异特征及分异规律,研究结果显示：（1）湘江流域上游传统村落平面基因变异相悖于本土的情况侧重发生在平面结构变异上;立面基因的变异相悖于本土、突变后直接消亡的情况较多;文化基因上受外来冲击的不适感较强,有加剧景观基因变异的风险和趋势。（2）湘江流域中游传统村落对立面基因的改动需求较大,且改动时间较早,造成立面基因变异以及平面基因中街巷格局、平面布局变异较大,变异后相悖于本土的特征显著;而在文化基因中,对文化的变异融合接受能力强,变异相融于本土的情况显著。（3）湘江流域下游传统村落在形态、布局特征明显的平面基因中产生的变异不大显著;在立面基因中遭遇突变后直接消亡的特征明显;在文化基因中,对家风信仰的基因变异以相融于本土为主,习俗礼仪的基因变异呈现分化趋势,单民族聚居式的家族以相悖于本土的特征为主,多民族聚居式的家族以相融于本土的特征为主。     

Clonal structure affects the assembling behavior in the Japanese queenless ant <Emphasis Type="Italic">Pristomyrmex punctatus</Emphasis>

The queenless ant Pristomyrmex punctatus (Hymenoptera: Myrmicinae) has a unique society that differs from those of other typical ants. This species does not have a queen, and the workers lay eggs and produce their clones parthenogenetically. However, a colony of these ants does not always comprise members derived from a single clonal line. In this study, we examined whether P. punctatus changes its “assembling behavior” based on colony genetic structure. We prepared two subcolonies—a larger one comprising 200 individuals and a smaller one comprising 100 individuals; these subcolonies were established from a single stock colony. We investigated whether these subcolonies assemble into a single nest. The genetically monomorphic subcolonies (single clonal line) always fused into a single nest; however, the genetically polymorphic subcolonies (multiple clonal lines) did not tend to form a single colony. The present study is the first to demonstrate that the colony genetic structure significantly affects social viscosity in social insects.