Solitary foundation or colony fission in ants: an intraspecific study shows that worker presence and number increase colony foundation success.

Dispersal and establishment strategies are highly variable. Each strategy is associated with specific costs and benefits, and understanding which factors favour or disfavour a strategy is a key issue in ecology and evolution. Ants exhibit several strategies of establishment, i.e. of colony foundation. Some species rely on winged queens that found new colonies alone when others found with accompanying workers (colony fission). The benefits conferred by these workers have been little studied and quantified, because comparing the costs and benefits of solitary foundation vs. colony fission is difficult when comparing different species. We investigated this using the ant Myrmecina graminicola, one of the few species that use both strategies. Young mated queens were allowed to found new colonies in the laboratory, with either zero (solitarily), two or four workers (colony fission). The presence of workers increased both survival and growth of the foundations over the first year, with more workers yielding higher growth. Few workers (as little as two workers) were sufficient to provide benefits, suggesting that in M. graminicola the strategy of colony fission may not dramatically decrease the number of new colonies produced compared to solitary foundation. Because queens performing solitary foundation or colony fission differ in dispersal (by flight vs. on foot), our results support the hypothesis that these two strategies of foundation coexist along a competition-colonization trade-off, where solitary foundation offers a colonization advantage, while colony fission has a competitive advantage.

Managing propagule pressure to prevent invasive species establishments: propagule size, number, and risk-release curve.

There is considerable evidence that keeping propagule pressure low can drastically reduce establishment probability of potential invasive species. Yet, most management plans and research efforts fail to explicitly acknowledge all three of the components of propagule pressure: size, number, and the risk-release relationship. It is unclear how failing to specify one or more of these components can influence the efficacy of management plans in preventing invasive species establishment. Furthermore, even if all components are acknowledged and quantified, there currently is no mathematical tool available to calculate the levels of propagule pressure that ensure attainment of a predetermined, and system-specific, target establishment probability. Here, we quantify the resulting uncertainty in establishment probability when one or more components of propagule pressure is unknown by using parameter uncertainty analysis on realistic values of propagule pressure. In addition, to aid in the development of management plans that explicitly set propagule pressure limits, we develop a propagule-pressure sensitivity analysis that we use to determine the required reduction in levels for propagule size and number (representative of management actions) to maintain a target establishment probability. We show that the precision of establishment estimates is highly dependent on knowledge of all three propagule pressure components, where the possible range of values for establishment probability can vary by over 50% without full specification. In addition, our sensitivity analysis showed that propagule size and number can be altered independently or in conjunction to lower establishment probability below a target level. Importantly, our sensitivity analysis was able to specifically quantify how much reduction in a propagule pressure component(s) is needed to reach a given target establishment probability. Our findings suggest that quantifying the three components of propagule pressure should be a priority for invasive species prevention moving forward. Furthermore, our sensitivity analysis tool can serve to guide the development of new invasive species management plans in a transparent and quantitative manner. Together with information on the costs associated with approaches to reducing propagule pressure, our tool can be used to identify the most cost-effective approach to prevent invasive species establishments.

Body size determines soil community assembly in a tropical forest.

Tropical forests shelter an unparalleled biological diversity. The relative influence of environmental selection (i.e., abiotic conditions, biotic interactions) and stochastic-distance-dependent neutral processes (i.e., demography, dispersal) in shaping communities has been extensively studied for various organisms, but has rarely been explored across a large range of body sizes, in particular in soil environments. We built a detailed census of the whole soil biota in a 12-ha tropical forest plot using soil DNA metabarcoding. We show that the distribution of 19 taxonomic groups (ranging from microbes to mesofauna) is primarily stochastic, suggesting that neutral processes are prominent drivers of the assembly of these communities at this scale. We also identify aluminium, topography and plant species identity as weak, yet significant drivers of soil richness and community composition of bacteria, protists and to a lesser extent fungi. Finally, we show that body size, which determines the scale at which an organism perceives its environment, predicted the community assembly across taxonomic groups, with soil mesofauna assemblages being more stochastic than microbial ones. These results suggest that the relative contribution of neutral processes and environmental selection to community assembly directly depends on body size. Body size is hence an important determinant of community assembly rules at the scale of the ecological community in tropical soils and should be accounted for in spatial models of tropical soil food webs.

Dynamics of Reintroduced Populations of Oedipoda caerulescens (Orthoptera, Acrididae) over 21 Years.

Conservation programs increasingly involve the reintroduction of animals which otherwise would not recolonize restored habitats. We assessed the long-term success of a project in which the Blue-winged grasshopper, Oedipoda caerulescens (L., 1758), was reintroduced to a nature reserve in Northwestern Switzerland, an alluvial gravel area where the species went extinct in the 1960s. In summer 1995, we released 110 individuals (50 females and 60 males) and 204 individuals (101 females and 103 males) into two restored gravel patches with sparse vegetation. We used a transect count technique to assess the population size of O. caerulescens in the years 1995-2004 and 2015-2016 and recorded the area occupied by the species. At both release sites, the populations persisted and increased significantly in size. Individuals that followed a newly created corridor established four new subpopulations. Seven years after reintroduction, O. caerulescens had reached a high abundance around the release sites and in the four colonized patches, indicating a successful project. At the same time, the dispersal corridor became increasingly overgrown by dense vegetation. Surveys 20 and 21 yr after introduction showed that the abundance of the Blue-winged grasshopper had strongly declined in the established subpopulations and moderately in the original release sites, owing to natural succession of the habitat and lack of disturbances, which reduced the area suitable for the species by 59%. Our study shows that reintroductions are unlikely to succeed without integration of long-term habitat management (in the present case maintenance of open ground).

Maternal body condition influences magnitude of anti-predator response in offspring.

Organisms exhibit plasticity in response to their environment, but there is large variation even within populations in the expression and magnitude of response. Maternal influence alters offspring survival through size advantages in growth and development. However, the relationship between maternal influence and variation in plasticity in response to predation risk is unknown. We hypothesized that variation in the magnitude of plastic responses between families is at least partly due to maternal provisioning and examined the relationship between maternal condition, egg provisioning and magnitude of plastic response to perceived predation risk (by dragonfly larvae: Aeshna spp.) in northern leopard frogs (Lithobates pipiens). Females in better body condition tended to lay more (clutch size) larger (egg diameter) eggs. Tadpoles responded to predation risk by increasing relative tail depth (morphology) and decreasing activity (behaviour). We found a positive relationship between morphological effect size and maternal condition, but no relationship between behavioural effect size and maternal condition. These novel findings suggest that limitations imposed by maternal condition can constrain phenotypic variation, ultimately influencing the capacity of populations to respond to environmental change.

Structured and unstructured intraspecific propagule trait variation across environmental gradients in a widespread mangrove.

Increasing studies have shown the importance of intraspecific trait variation (ITV) on ecological processes. However, the patterns and sources of ITV are still unclear, especially in the propagules of coastal vegetation. Here, we measured six hypocotyl traits for 66 genealogies of Kandelia obovata from 26 sites and analyzed how ITV in these traits was distributed across geography and genealogy through variance partitioning. We further constructed mixed models and structural equation models to disentangle the effects of climatic, oceanic, and maternal factors on ITV. Results showed that size-related traits decreased along increasing latitudinal gradients, which was mainly driven by positive regulation of temperature on these traits. By contrast, ITV of shape trait was unstructured along latitudinal gradients and did not show any dependence among environmental variables. These findings indicate that propagule size mainly varied between populations, whereas propagule shape mainly varied between individuals. Our study may provide useful insights into the ITV in propagule from different functional dimensions and on a broad scale, which may facilitate mangrove protection in light of ITV.

Time-lag in extinction dynamics in experimental populations: evidence for a genetic Allee effect?

1. Propagule pressure, i.e. the number of individuals introduced, is thought to be a major predictor of the establishment success of introduced populations in the field. Its influence in laboratory experimental systems has however been questioned. In fact, other factors involved in long-term population persistence, like habitat size, were usually found to explain most of the dynamics of experimental populations. 2. To better understand the respective influence of short- and long-term factors and their potential interaction on extinction dynamics in experimental systems, we investigated the influence of propagule pressure, habitat size and genetic background on the early dynamics of laboratory-based populations of a hymenopteran parasitoid. 3. The amount of demographic variance differed between establishment and persistence phase and was influenced by habitat size and genetic background (geographic strain), but independent of propagule pressure. In contrast, the probability of extinction within five generations depended on the genetic background and on the interaction between propagule pressure and habitat size. Vulnerability to extinction in small size habitats was increased when populations were founded with a small number of individuals, but this effect was delayed until the third to fifth generations. 4. These results indicate that demographic stochasticity is influential during population establishment, but is not affected by the genetic variability of propagules. On the other hand, extinction might be influenced by a genetic Allee effect triggered by the combination of low propagule pressure and genetic drift. Finally, we documented consistent differences between genetic backgrounds in both deterministic and stochastic population dynamics patterns, with major consequences on extinction risk and ultimately population establishment.

The positive correlation between maternal size and offspring size: fitting pieces of a life-history puzzle.

The evolution of investment per offspring (I) is often viewed through the lens of the classic theory, in which variation among individuals in a population is not expected. A substantial departure from this prediction arises in the form of correlations between maternal body size and I, which are observed within populations in virtually all taxonomic groups. Based on the generality of this observation, we suggest it is caused by a common underlying mechanism. We pursue a unifying explanation for this pattern by reviewing all theoretical models that attempt to explain it. We assess the generality of the mechanism upon which each model is based, and the extent to which data support its predictions. Two classes of adaptive models are identified: models that assume that the correlation arises from maternal influences on the relationship between I and offspring fitness [w(I)], and those that assume that maternal size influences the relationship between I and maternal fitness [W(I)]. The weight of evidence suggests that maternal influences on w(I) are probably not very general, and even for taxa where maternal influences on w(I) are likely, experiments fail to support model predictions. Models that assume that W(I) varies with maternal size appear to offer more generality, but the current challenge is to identify a specific and general mechanism upon which W(I) varies predictably with maternal size. Recent theory suggests the exciting possibility that a yet unknown mechanism modifies the offspring size-number trade-off function in a manner that is predictable with respect to maternal size, such that W(I) varies with size. We identify two promising avenues of inquiry. First, the trade-off might be modified by energetic costs that are associated with the initiation of reproduction ('overhead costs') and that scale with I, and future work could investigate what specific overhead costs are generally associated with reproduction and whether these costs scale with I. Second, the trade-off might be modified by virtue of condition-dependent offspring provisioning coupled with metabolic factors, and future work could investigate the proximate cause of, and generality of, condition-dependent offspring provisioning. Finally, drawing on the existing literature, we suggest that maternal size per se is not causatively related to variation in I, and the mechanism involved in the correlation is instead linked to maternal nutritional status or maternal condition, which is usually correlated with maternal size. Using manipulative experiments to elucidate why females with high nutritional status typically produce large offspring might help explain what specific mechanism underlies the maternal-size correlation.

Adaptive plasticity of egg size in response to competition in the cowpea weevil, Callosobruchus maculatus (Coleoptera: Bruchidae).

Life history theory predicts that larger propagules should be produced when the offspring are expected to experience intense competition. This study tested whether female cowpea weevils responded to high larval or adult density by producing larger eggs. In a splitbrood design I measured the effect of density experienced by females at their larval stage (1 vs. 4-6 larvae/cowpea) on the size of eggs produced just after emergence. The females were then kept either at low adult density (1 female+1 male per vial), or at high adult density (10 females+10 males) for 2 days, and tested for the effect of this adult density treatment on the size of eggs laid subsequently. I measured egg length and width, as well as the diameter of the entrance tunnel made by the larva, which can be regarded as a crude measure of larval size. Females that experienced high adult density subsequently laid slightly wider eggs than those kept at low density. This difference, albeit small (about 1-4% after correction for female weight and the effect of family, depending on the statistical model used), was statistically significant and robust to alterations of the statistical model. It may be a remnant of a larger plastic response of egg size to competition that has become eroded during many generations in high-density laboratory cultures. There was no difference in egg length or the diameter of the entrance tunnel. Eggs laid just after emergence by females reared at high larval density also tended to be wider than those produced by females that had no competitors. This effect was only marginally significant, however, and sensitive to the statistical model. Both egg length and width and the diameter of the entrance tunnel increased with female weight and decreased with female age. The tunnel diameter was positively correlated with both egg length and width, but the effect of width was larger.

Effects of emergence date and maternal size on egg development and sizes of eggs and first-instar nymphs of a semelparous aquatic insect.

We examined whether or not sizes of eggs and offspring were related to emergence date or maternal size in a semelparous aquatic insect (the burrowing mayfly, Hexagenia) in which parental care is lacking and oviposited eggs are passively dispersed. We quantified the size of males and female imagos over the emergence span at a site on the Detroit River, Canada, and investigated relationships between emergence date and female size and (1) egg size and (2) size of first-instar nymphs. Although size of female imagos (H. limbata and H. rigida combined) declined significantly (P<0.025) over the emergence season, there was no significant relationship between body length and emergence date for males of either species. Males were significantly (P<0.001) smaller than females. H. limbata eggs, subsampled from three individuals from each of three size classes of female imagos collected on seven sampling dates, were measured using video image analysis. Eggs (n=100) oviposited by each of 63 H. limbata imagos were inspected daily for hatching. Newly hatched nymphs were removed, counted and measured. Egg size (P<0.001) and size of first-instar nymphs (P<0.001) varied significantly with emergence date, but not maternal size. The largest eggs and newly hatched nymphs occurred at peak emergence of adults. The synchronous release of larger (faster-sinking) eggs may result in reduced predation. Plasticity in egg development time and egg and nymph size may account for the ability of this taxon to recover from episodes of massive population reduction.

The evolution of offspring size and number: a test of the Smith-Fretwell model in three species of crickets.

Most models of parental investment in offspring assume a trade-off between propagule size and number, and an increasing "concave down" function relating offspring fitness to propagule size. In this study, we test these two fundamental assumptions, using three closely related species of crickets, Gryllus firmus, G. veletis, and G. pennsylvanicus. Egg weight, 35-day fecundity and 35-day egg biomass were estimated in a population of each species, and the relationships between these reproductive traits and date of egg laying and body size were estimated. The relationships between egg weight and offspring survival were also sought for eggs buried at different depths, soil moistures, and soil types (G. firmus and G. veletis), as well as in the field (G. pennsylvanicus). A trade-off between egg weight and 35-day fecundity was revealed in a multivariate analysis taking into account among-species variation in egg weight and body size. Independent of the environmental conditions affecting the eggs, a positive correlation existed between the number of larvae that emerged from the soil and propagule weight in each species. Therefore, these findings provide partial support for the assumptions considered in the models mentioned above. A single optimal egg size was favored in two out of the three sets of conditions in which the functions relating egg weight to larval survival could be derived. The conditions encountered by the eggs, however, influenced the average survival of the larvae, as well as the shape of the relationship between egg weight and offspring survival. This suggests that cricket eggs frequently face heterogeneous environments with respect to egg and hatchling survival; the implication of habitat heterogeneity on the evolution of an optimal egg size is considered. The relationships between the reproductive components and female age and size, as well as between egg size and variation in cricket life-history, are discussed in an ecological and evolutionary context.