The predictability of evolution: glimpses into a post-Darwinian world

The very success of the Darwinian explanation, in not only demonstrating evolution from multiple lines of evidence but also in providing some plausible explanations, paradoxically seems to have served to have stifled explorations into other areas of investigation. The fact of evolution is now almost universally yoked to the assumption that its outcomes are random, trends are little more than drunkard’s walks, and most evolutionary products are masterpieces of improvisation and far from perfect. But is this correct? Let us consider some alternatives. Is there evidence that evolution could in anyway be predictable? Can we identify alternative forms of biological organizations and if so how viable are they? Why are some molecules so extraordinarily versatile, while others can be spoken of as “molecules of choice”? How fortuitous are the major transitions in the history of life? What implications might this have for the Tree of Life? To what extent is evolutionary diversification constrained or facilitated by prior states? Are evolutionary outcomes merely sufficient or alternatively are they highly efficient, even superb? Here I argue that in sharp contradistinction to an orthodox Darwinian view, not only is evolution much more predictable than generally assumed but also investigation of its organizational substrates, including those of sensory systems, which indicates that it is possible to identify a predictability to the process and outcomes of evolution. If correct, the implications may be of some significance, not least in separating the unexceptional Darwinian mechanisms from underlying organizational principles, which may indicate evolutionary inevitabilities.     

Multilevel selection and social evolution of insect societies

How sterile, altruistic worker castes have evolved in social insects and how they are maintained have long been central topics in evolutionary biology. With the advance of kin selection theory, insect societies, in particular those of haplodiploid bees, ants, and wasps, have become highly suitable model systems for investigating the details of social evolution and recently also how within-group conflicts are resolved. Because insect societies typically do not consist of clones, conflicts among nestmates arise, for example about the partitioning of reproduction and the allocation of resources towards male and female sexuals. Variation in relatedness among group members therefore appears to have a profound influence on the social structure of groups. However, insect societies appear to be remarkably robust against such variation: division of labor and task allocation are often organized in more or less the same way in societies with high as in those with very low nestmate relatedness. To explain the discrepancy between predictions from kin structure and empirical data, it was suggested that constraints—such as the lack of power or information—prevent individuals from pursuing their own selfish interests. Applying a multilevel selection approach shows that these constraints are in fact group-level adaptation preventing or resolving intracolonial conflict. The mechanisms of conflict resolution in insect societies are similar to those at other levels in the biological hierarchy (e.g., in the genome or multicellular organisms): alignment of interests, fair lottery, and social control. Insect societies can thus be regarded as a level of selection with novelties that provide benefits beyond the scope of a solitary life. Therefore, relatedness is less important for the maintenance of insect societies, although it played a fundamental role in their evolution.Dedicated to Prof. Ernst Mayr     

Pre- and post-hatch trophic egg production in the subsocial burrower bug, Canthophorus niveimarginatus (Heteroptera: Cydnidae)

In recent years, three terrestrial bugs, Adomerus triguttulus and Sehirus cinctus (Cydnidae) and the closely related Parastrachia japonensis (Parastrachiidae), have been the focus of several fascinating studies because of the remarkable, extensive parental care they were found to display. This care includes egg and nymph guarding, production of trophic eggs, unfertilized, low cost eggs that are used as food by newly hatched nymphs, and progressive provisioning of the host seed. In this study, we have investigated yet a third related Asian cydnid, Canthophorus niveimarginatus, with regard to the possible occurrence of some or all of these complex traits in order to assess how widespread these maternal investment patterns are in this group of insects and to better understand the implications of their manifestations from an evolutionary context. Manipulative experiments were carried out in the lab to determine whether females provision nests. Observational and egg removal studies were carried out to determine whether trophic eggs are produced, and, if they are, their possible impact on nymphal success. The findings revealed that C. niveimarginatus does, in fact, progressively provision young, and this species also displays all of the other behaviors associated with extended parental care in subsocial insects. Moreover, unlike the other two related species, which place trophic eggs on the surface of the original egg mass, C. niveimarginatus produces both pre- and post-hatch trophic eggs. Nymphs deprived of access to post-hatch trophic eggs had significantly lower body weight and survival rate than those that fed on them. To our knowledge, this is the first time the production of both pre- and post-hatch trophic eggs has been demonstrated in insects outside the Hymenoptera. In this paper, we qualitatively and quantitatively demonstrate the provisioning behavior and patterns of trophic egg production in C. niveimarginatus. When and how trophic eggs are produced and delivered to young should have important correlations with the ecological and life history constraints under which a species has evolved. Thus, we also discuss the possible ecological and life history factors that favor the evolution of post-hatch trophic eggs.     

Moorhens have an internal representation of their own eggs

How do birds recognize their own eggs? Do they have a stored template for their own egg characteristics, or do they use another mechanism? Intraspecific brood parasitism is considered to be an additional reproductive tactic where females can increase their own reproductive success. Because of the costs involved in rearing young that are not their own, it will pay females to detect and reject the eggs of a parasite, although it is not known how they do this. Here, we show experimentally that moorhens will cease laying in a nest when their first egg is replaced with another hen’s egg but not when it is replaced with their own egg taken from an earlier clutch. This provides good evidence that birds have an internal representation of their own eggs and use this in decisions about whether to reject foreign eggs.     

Predicting fish population dynamics: compensation and the importance of site-specific considerations

The magnitude of the compensatory response inherent to fish population seems to be questioned with each new generation of scientists and regulators. The major reason for this is the demand by some that compensation be completely understood at the process level before compensation can be factored into population predictions. We first discuss several lines of evidence for the existence of compensation, and the reasons why compensation is so difficult to measure at the process level. We then offer life history theory as a general framework for improving our understanding of the likely magnitude of compensation in fish population dynamics. We use detailed, individual-based models and field data for yellow perch-walleye, striped bass, and a lake community to illustrate the complexity of compensation, and to show that predictions for a particular population require site-specific information. Analyses show that the life stage and magnitude of density-dependence can vary for different populations of the same species, and that population responses can depend on the arrangement of the food web. We must stop reinventing the density-dependence wheel; too many resources (both natural and monetary) are at stake. We suggest using three approaches for addressing the compensation issue: (1) life history theory for providing a sound basis for qualitatively understanding fish population dynamics and the magnitude of compensation; (2) age- and stage-structured matrix models for screening-level analyses to identify populations that may be at unacceptable risks of decline and therefore require additional analyses; and (3) individual-based models for better understanding the complexities of compensation at the process level and for impact assessment of at-risk, well-studied populations.     

Interphyletic relationships in the use of nesting cavities: mutualism,competition and amensalism among hymenopterans and vertebrates

Although competition is usually assumed to be the most common interaction between closely related organisms that share limiting resources, the relationships linking distant taxa that use the same nesting sites are poorly understood. In the present study, we examine the interactions among social hymenopterans (honeybees and wasps) and vertebrates in tropical ecosystems of East Africa. By analysing the preferences of these three groups for nest boxes that were empty or previously occupied by a different taxon, we try to establish whether the relationships among them are commensal, mutualistic, competitive or amensal. Vertebrates and honeybees selected nest boxes that had previously been occupied by the other, which suggests that each obtains some benefit from the other. This relationship can be considered mutualistic, although a mutual preference for each others’ nests does not exclude a competitive interaction. Vertebrates and wasps preferred nest boxes not previously occupied by the other, which suggests that they compete for tree cavities. Finally, wasps seemed to completely refuse cavities previously used by honeybees, while the bees occupied cavities regardless of whether they had been previously used by wasps, an apparently amensal relationship. These results indicate that the interdependence between distantly related taxa is stronger and more complex than previously described, which may have important implications for population dynamics and community structure.

     

Governance of complex environmental situations through social learning: a synthesis of SLIM's lessons for research,policy and practice

This synthesis of the SLIM project findings deals with the development and deployment of knowledge and research that is useful for actions that transform at socially and ecologically meaningful scales. A diagnostic framework (DF) is elaborated that aims to transform the findings into a tool that could bring stakeholders, in other contexts, to understand better their own roles in complex natural resource management situations. The DF invites the user to engage in successive stages of comprehension by exploring: what are complex situations of change about? What are the main components involved? Why are these components important? How do they influence what we know and how we act? What could be our role in changing the situation? We identify five ‘variables’ that together can account systemically for transformation processes that are constituted in social learning and concerted actions. We show how the DF may be used in situations of complexity and uncertainty by researchers, acting variously as observer, facilitator or co-researcher, and how it may help to guide research practice. We conclude by consolidating key messages about the relationship between knowledge, research, and policy and the main implications for water managers of being open to social learning processes.     

经济集聚对中国三大城市群土地利用生态效率的影响

基于非期望产出的超效率EBM模型,测算2005—2016年京津冀、长三角和珠三角等中国三大城市群土地利用生态效率,采用核密度函数对其演变特征进行分析,并运用面板门槛回归模型和面板向量自回归模型,从产业和人口两个角度分析经济集聚对城市群土地利用生态效率的非线性影响和直接冲击。结果表明：（1）2005—2016年间中国三大城市群土地利用生态效率总体呈现先收敛后分化的演化特征,不同城市群之间的土地利用生态效率时空演变具有显著差异;（2）第三产业集聚水平和人口集聚水平对城市群土地利用生态效率表现出单门槛效应,表明适度产业集聚和人口集聚对城市群土地利用生态效率具有正向促进作用,反之则会导致促进作用下降甚至出现负向影响;（3）第三产业集聚水平和土地利用生态效率自身对土地利用生态效率的直接冲击都是从开始就达到顶峰而后收敛为0,而第二产业集聚水平和人口集聚水平则是先负向冲击后持续正向冲击。城市群在制定经济发展政策和土地利用政策时,既要充分发挥经济集聚的正面环境效应,也要重视不同经济集聚的适度性和动态性。     

Seasonality of spawning by tropical anguillid eels around Sulawesi Island, Indonesia

Remarkably little is known about the life histories of the many tropical anguillid eels distributed across the Indo-Pacific region, and since the Danish expedition to study eels in the region in 1928 and 1929, research on these eels has only begun again in recent years. Sampling for anguillid leptocephali in the Indonesian Seas has been carried out recently to learn about the spawning ecology and larval distributions of tropical eels there. The leptocephali of Anguilla marmorata, Anguilla bicolor pacifica, Anguilla borneensis, Anguilla interioris, and Anguilla celebesensis were collected around Sulawesi Island both in May 2001 and October of 2002. The development of genetic identification techniques has enabled these leptocephali to be identified to species, and their distributions and sizes during different seasons indicated that there are differing life history patterns among sympatric species in the region. A. celebesensis was found to have been spawning in Tomini Bay of northeastern Sulawesi Island in March and April 2001, but apparently, no spawning had occurred in the late summer and fall of 2002. Studies on anguillid glass eels have suggested that tropical anguillids may spawn throughout much of year, but our research on leptocephali in Tomini Bay and data on the downstream migration of tropical anguillids in the major tributary to Tomini Bay indicate that A. celebesensis may have a distinct seasonal pattern of spawning possibly related to the regional monsoon cycles. This is the first evidence of seasonality of spawning in tropical anguillid eels whose life histories are only just beginning to be revealed.     

Climatic influences on the breeding biology of the agile frog (<Emphasis Type="Italic">Rana dalmatina</Emphasis>)

Severe population declines of amphibians have been shown to be attributed to climate change. Nevertheless, the various mechanisms through which climate can influence population dynamics of amphibians remain to be assessed, notably to disentangle the relative synergetic or antagonistic influences of temperature and precipitations on specific life history stages. We investigated the impact of rainfall and temperature on the egg-clutch abundance in a population of agile frog (Rana dalmatina) during 29 years (1987–2016) on 14 breeding sites located in Brenne Natural Park, France. Specifically, we examined the influence of environmental conditions occurring during five temporal windows of the year cycle corresponding to specific life history stages. Overall, our results suggest that the year-to-year fluctuations of egg-clutch abundances in Brenne Natural Park were partly dependent on local climatic conditions (rainfall and temperature). Climate seemed to influence breeding frogs during the autumn-winter period preceding reproduction. Spring and summer conditions did not influence reproduction. Additionally, we failed to detect effects of climatic conditions on newly metamorphosed individuals. Other factors such as density dependence and inter-specific interactions with introduced predators are likely to play a significant role in reproduction dynamics of the studied frog populations.     

Plant invasion phenomenon enhances reproduction performance in an endangered spider

Current models in evolutionary ecology predict life history alterations in response to habitat suitability to optimize fitness. Only few empirical studies have demonstrated how life history traits that are expected to trade off against each other differ among environments. In Europe, many salt marshes have been recently invaded by the grass Elymus athericus. Previous studies however showed higher densities of the endangered spider Arctosa fulvolineata (Araneae: Lycosidae) in invaded salt marshes compared to natural habitats, which suggests a lower habitat suitability in the latter. The aim of this study was to determine if this emerging habitat (1) affects the amount of resource acquisition and (2) alters the balance between life history traits that are expected to trade off against each other in this stenotopic salt marsh species. As suggested by theoretical studies, an optimization of fitness by increasing egg size at the cost of decreasing fecundity in unsuitable (i.e., natural) habitats was expected. Females presenting cocoon were then collected in close invaded and natural salt marsh areas within the Mont Saint-Michel Bay (France). By considering female mass as covariate, cocoon mass, number of eggs, and egg volume were compared between both habitats. Clutch mass was strongly determined by female mass in both habitats. Clutch mass was however significantly smaller in the natural habitat compared to the invaded habitat, indicating a higher resource acquisition in the latter. When correcting for female size, fecundity was additionally increased in the invaded habitat through a significant decrease in egg size. This phenotypic response can be explained by differences in habitat structure between invaded and natural habitats: the former offers a more complex litter favoring nocturnal wanderers like A. fulvolineata. The existence of such an adaptive reproduction strategy depending on habitat suitability constitutes an original case of an invasion that favors an endangered species.     

Amphimixis and the individual in evolving populations: does Weismann’s Doctrine apply to all,most or a few organisms?

The German biologist August Weismann (1834–1914) proposed that amphimixis (sexual reproduction) creates variability for natural selection to act upon, and hence he became one of the founders of the Neo-Darwinian theory of biological evolution. He is perhaps best known for what is called “Weismann’s Doctrine” or “Weismann’s Barrier” (i.e. the irreversible separation of somatic and germ cell functionalities early during ontogeny in multicellular organisms). This concept provided an unassailable argument against “soft inheritance” sensu Lamarck and informed subsequent theorists that the only “individual” in the context of evolution is the mature, reproductive organism. Herein, we review representative model organisms whose embryology conforms to Weismann’s Doctrine (e.g. flies and mammals) and those that do not (e.g. freshwater hydroids and plants) based on this survey and the Five Kingdoms of Life scheme; we point out that most species (notably bacteria, fungi, protists and plants) are “non-Weismannian” in ways that make a canonical definition of the “individual” problematic if not impossible. We also review critical life history functional traits that allow us to create a matrix of all theoretically conceivable life cycles (for eukaryotic algae, embryophytes, fungi and animals), which permits us to establish where this scheme Weismann’s Doctrine holds true and where it does not. In addition, we argue that bacteria, the dominant organisms of the biosphere, exist in super-cellular biofilms but rarely as single (planktonic) microbes. Our analysis attempts to show that competition among genomic variants in cell lineages played a critical part in the evolution of multicellularity and life cycle diversity. This feature was largely ignored during the formulation of the synthetic theory of biological evolution and its subsequent elaborations.     

Social Rank, Stress, Fitness, and Life Expectancy in Wild Rabbits

 Wild rabbits of the two sexes have separate linear rank orders, which are established and maintained by intensive fights. The social rank of individuals strongly influence their fitness: males and females that gain a high social rank, at least at the outset of their second breeding season, have a much higher lifetime fitness than subordinate individuals. This is because of two separate factors: a much higher fecundity and annual reproductive success and a 50% longer reproductive life span. These results are in contrast to the view in evolutionary biology that current reproduction can be increased only at the expense of future survival and/or fecundity. These concepts entail higher physiological costs in high-ranking mammals, which is not supported by our data: In wild rabbits the physiological costs of social positions are caused predominantly by differential psychosocial stress responses that are much lower in high-ranking than in low-ranking individuals. Received: 23 February 1999 / Accepted: 17 March 1999     

Application of a population dynamics model to the probabilistic assessment of cooling water intake effects of Millstone Nuclear Power Station (Waterford,CT) on a nearby winter flounder spawning stock

A major concern of the Connecticut Department of Environmental Protection at Millstone Station has been the long-term effects of the station’s Cooling Water Intakes System (CWIS) on a small stock of winter flounder spawning in the Niantic River. Following the selection of a population modeling approach for long-term impact assessment in 1983, sampling programs were directed to collect data specifically for this purpose. These data were used to estimate the size of the local flounder stock and the fraction of its annual production lost to CWIS effects, and also evaluate a possible relationship between late-winter water temperatures and recruitment success of flounder. Since the mid 1980s, the main assessment tool at Millstone has been an age-structured population model that incorporated uncertainty in selected model parameters and provided output suitable for probabilistic analyses of simulation results. This model projected annual population sizes resulting from different fishing rates, larval entrainment losses, and impingement of juvenile and adult fish. Compensation was introduced via a Ricker stock-recruit function with an additional term to describe the effect of water temperature on recruitment success. Ricker’s α parameter, which describes the species inherent capacity to increase in numbers, was derived by four indirect methods based on life history parameters. In the basic simulation strategy, an unfished flounder stock was simulated first, and then fishing effects and mortality of the young attributed to CWIS effects were added. Combinations of different fishing rates and CWIS effects, either estimated or hypothesized, could be simulated for comparing the effectiveness of different mitigation alternatives. This approach helped identify fishing mortality as the driving force shaping the future size and viability of the local flounder population at Millstone and suggested that larval entrainment mitigation measures, costing hundreds of millions of dollars, would be ineffective in reversing currently declining trends of that population.     

Induced resistance enzymes in wild plants–do ‘early birds’ escape from pathogen attack?

Systemic acquired resistance (SAR) of plants to pathogens is a well-defined phenomenon. The underlying signalling pathways and its application in crop protection are intensively studied. However, most studies are conducted on crop plants or on Arabidopsis as a model plant. The taxonomic distribution of this phenomenon and its dependence on life history are thus largely unknown. We quantified activities of three classes of resistance-related enzymes in 18 plant species to investigate whether plants with varying life histories differ in their investment in disease resistance. Enzyme activities were quantified in untreated plants, and in plants induced with BION, a chemical resistance elicitor. All species showed constitutive activities of chitinase, peroxidase, or glucanase. However, constitutive chitinase activities varied by 30 times, and peroxidase by 50 times, among species. Several species did not respond to the induction treatment, while enzyme activities in other species increased more than threefold after BION application. Plant species differ dramatically in the presence and inducibility of resistance enzymes. This variation could be related to life history: While all resistance enzymes were significantly induced in larger perennial plants that flower during summer, spring geophytes hardly showed inducible resistance. These plants grow in an environment that is characterised by a low-pathogen pressure, and thus may simply ‘escape’ from infection. Our study presents the first comparative data set on resistance-related enzymes in noncultivated plants. The current view on SAR—narrowed by the concentration on cultivated crops—is not sufficient to understand the ecological and evolutionary relevance of this widespread plant trait.     

The water-energy-food nexus: Trade-offs,thresholds and transdisciplinary approaches to sustainable development

Global challenges have exacerbated a search for solutions to poverty and environmental degradation. Integration it was argued would help address the twin challenge. Integrated Water Resources Management (IWRM) was supposed to be that magic bullet and was embraced by scientists because of the clinical efficiency with which it argued for integrated analysis of sectors and resources and of systems and scale conditions. This paper argues that effective implementation of the Water-Energy-Food (WEF) Nexus can be supported by robust science. The corollary that robust science automatically leads to effective implementation is not always known to be true. The nexus approach sheds light on the challenges of implementation by introducing concepts of trade-offs and thresholds and consequently emphasizes the importance of transdisciplinary approaches to sustainable development. This paper reviews the results of recent research to offer tentative answers to the following questions: (a) Why is the governance dimension important to undertake an integrated analysis of water-energy-food challenges? (b) What does the nexus approach connote in normative and institutional terms? (c) What does implementation mean in nexus terms? (d) How can we establish if the nexus approach is an improvement over business as usual? and (e) What tools are available that would enable translation of results of scientific research to create an evidence base that would enable decision makers to act in support of sustainable development?     

Geographic variations of life history traits and potential trade-offs in different populations of the parasitoid Leptopilina heterotoma

Energy allocation is determined by resource availability and trade-offs among traits, and so organisms have to give some traits priority over others to maximize their fitness according to their environment. In this study, we investigated the geographic variations in life history traits and potential trade-offs in populations of the parasitoid Leptopilina heterotoma (Hymenoptera: Figitidae) originating from the north and the south of the Rh?ne–Sa?ne valley (over a gradient of 300?km, South-East France). We measured a set of traits related to reproduction, maintenance, and mobility using several estimators of each of these main functions determined at different times. We did not find any clear differences between populations from contrasting areas, whereas the southern populations, which were all assumed to be exposed to similar environmental conditions, displayed contrasting patterns of energy allocation. Thus, the most likely explanation seems to be that the evolution of the life history of L. heterotoma is probably shaped by local selective pressures, such as microclimate, microhabitats, or intensity of competition, rather than by regional ecological conditions. Using our study as an example, we discuss the interest of considering several traits and using different ways of measuring them, concluding that multiple measurements should be performed in future studies to ensure the robustness of the results.     

Symbiosis and Pathogenesis: Evolution of the Microbe-Host Interaction

Symbiotic and pathogenic bacteria have in common that they live in or on host organisms or host cells. To make a successful living in eukaryotic hosts, bacteria must possess the traits to recognize a given host and establish adherence. When the bacterial location is internal or intracellular, they must further have the ability to invade, to establish a niche, and finally to multiply within a host. The underlying mechanisms which allow this form of existence show similarities between symbiotic and pathogenic bacteria. The final outcome, however, may result in a wide spectrum of consequences for the host ranging from the acquisition of novel metabolic pathways to damage or death. Despite the vastly different forms of interactions, symbiotic and pathogenic bacteria have in common that they are adapted to a particular environmental niche represented by the host organism or compartment thereof. This contribution reviews the evolutionary forces which have shaped the microbial-host interactions. Particular emphasis is placed on the genetic and molecular mechanisms that drive bacterial evolution in response to the selective pressures of the host environment.