Hierarchies and the Sloshing Bucket: Toward the Unification of Evolutionary Biology

Evolutionary biology presents a bewildering array of phenomena to scientists and students alike—ranging from molecules to species and ecosystems; and embracing 3.8 billion years of life’s history on earth. Biological systems are arranged hierarchically, with smaller units forming the components of larger systems. The evolutionary hierarchy, based on replication of genetic information and reproduction, is a complex of genes/organisms/demes/species and higher taxa. The ecological hierarchy, based on patterns of matter–energy transfer, is a complex of proteins/organisms/avatars/local ecosystems/regional ecosystems. All organisms are simultaneously parts of both hierarchical systems. Darwin’s original formulation of natural selection maps smoothly onto a diagram where the two hierarchical systems are placed side-by-side. The “sloshing bucket” theory of evolution emerges from empirical cases in biological history mapped onto this dual hierarchy scheme: little phenotypically discernible evolution occurs with minor ecological disturbance; conversely, greatest concentrations of change in evolutionary history follow mass extinctions, themselves based on physical perturbations of global extent. Most evolution occurs in intermediate-level regional “turnovers,” when species extinction leads to rapid evolution of new species. Hierarchy theory provides a way of integrating all fields of evolutionary biology into an easily understood—and taught—rubric.

Biological species is the only possible form of existence for higher organisms: the evolutionary meaning of sexual reproduction

   

Consistent holistic view of sexual species as the highest form of biological existence is presented. The Weismann's idea that sex and recombination provide the variation for the natural selection to act upon is dominated in most discussions of the biological meaning of the sexual reproduction. Here, the idea is substantiated that the main advantage of sex is the opposite: the ability to counteract not only extinction but further evolution as well. Living systems live long owing to their ability to reproduce themselves with a high fidelity. Simple organisms (like bacteria) reach the continued existence due to the high fidelity of individual genome replication. In organisms with a large genome and complex development, the achievable fidelity of DNA replication is not enough for the precise reproduction of the genome. Such species must be capable of surviving and must remain unchanged in spite of the continuous changes of their genes. This problem has no solution in the frame of asexual ("homeogenomic") lineages. They would rapidly degrade and become extinct or blurred out in the course of the reckless evolution. The core outcome of the transition to sexual reproduction was the creation of multiorganismic entity - biological species. Individual organisms forfeited their ability to reproduce autonomously. It implies that individual organisms forfeited their ability to substantive evolution. They evolve as a part of the biological species. In case of obligatory sexuality, there is no such a thing as synchronic multi-level selection. Natural selection cannot select anything that is not a unit of reproduction. Hierarchy in biology implies the functional predestination of the parts for the sake of the whole. A crucial feature of the sexual reproduction is the formation of genomes of individual organisms by random picking them over from the continuously shuffled gene pool instead of the direct replication of the ancestor's genome. A clear anti-evolutionary consequence of the sexuality is evident from the fact that the genotypes of the individuals with an enhanced competitiveness are not transmitted to the next generation. Instead, after mating with "ordinary" individuals, these genotypes scatter and rearrange in new gene combinations, thus preventing the winner from exploiting the success.     

First Chemical Analysis and Characterization of the Male Species‐Specific Cephalic Labial‐Gland Secretions of South American Bumblebees

The evolution of signals and reproductive traits involved in the pre‐mating recognition has been in focus of abundant research in several model species, such as bumblebees (genus Bombus). However, the most‐studied bumblebee reproductive trait, the male cephalic labial gland secretions (CLGS), remains unknown among bumblebee species from South America. In this study, the CLGS of five South American bumblebees of the subgenera Thoracobombus (Bombus excellens and B. atratus) and Cullumanobombus (B. rubicundus, B. hortulanus, and B. melaleucus) were investigated, by comparing the chemical compositions of their secretions to those of closely related European species. The results showed an obvious interspecific differentiation in both subgenera. The interspecific differentiation among the species of the Thoracobombus subgenus involved different compounds present at high contents (main compounds), while those of the Cullumanobombus subgenus shared the same main components. This suggests that among the species of the Cullumanobombus subgenus, the differentiation in minor components could lead to species discrimination.     

Leaf and stem physiological responses to summer and winter extremes of woody species across temperate ecosystems

Winter cold limits temperate plant performance, as does summer water stress in drought‐prone ecosystems. The relative impact of seasonal extremes on plant performance has received considerable attention for individual systems. An integrated study compiling the existing literature was needed to identify overall trends. First, we conducted a meta‐analysis of the impacts of summer and winter on ecophysiology for three woody plant functional types (winter deciduous angiosperms, evergreen angiosperms and conifers), including data for 210 records from 75 studies of ecosystems with and without summer drought across the temperate zone. Second, we tested predictions by conducting a case study in a drought‐prone Mediterranean ecosystem subject to winter freezing. As indicators of physiological response of leaves and xylem to seasonal stress, we focused on stomatal conductance (g_s), percent loss of stem xylem hydraulic conductivity (PLC) and photochemical efficiency of photosystem II (F_v/F_m). Our meta‐analysis showed that in ecosystems without summer drought, g_s was higher during summer than winter. By contrast, in drought‐prone ecosystems many species maintained open stomata during winter, with potential strong consequences for plant carbon gain over the year. Further, PLC tended to increase and F_v/F_m to decrease from summer to winter for most functional types and ecosystems due to low temperatures. Overall, deciduous angiosperms were most sensitive to climatic stress. Leaf gas exchange and stem xylem hydraulics showed a coordinated seasonal response at ecosystems without summer drought. In our Mediterranean site subjected to winter freezing the species showed similar responses to those typically found for ecosystems without summer drought. We conclude that winter stress is most extreme for systems without summer drought and systems with summer drought and winter freezing, and less extreme for drought‐prone systems without freezing. In all cases the evergreen species show less pronounced seasonal responses in both leaves and stems than deciduous species.     

Thoughts on the ecology and evolution of the intertidal biota of the Azores and other Atlantic islands

The special features of the intertidal ecosystems of remote islands are reviewed briefly before focusing on the littoral zone of Macaronesia (Azores, Madeira and Canaries). Distribution patterns are briefly compared with other European shores. Species missing from the Azores compared with Madeira, the Canaries and continental European shores are listed. The degree of genetic differentiation within and amongst selected important species of gastropods (Patella spp.) and barnacles (Chthamalus spp.) in Macaronesia is reviewed. The role of isolated islands in allopatric speciation of these organisms is discussed in relation to dispersal. Possible threats to the populations and communities of rocky shores in Macaronesia are considered in the context of the endemic nature of certain species. The importance of studies of basic ecology and genetics to inform resource management and conservation is highlighted. Future research directions are indicated, emphasizing the usefulness of Macaronesia as a model system to explore speciation in intertidal organisms.     

Trophic-dynamic considerations in relating species diversity to ecosystem resilience

Complexity in the networks of interactions among and between the living and abiotic components forming ecosystems confounds the ability of ecologists to predict the economic consequences of perturbations such as species deletions in nature. Such uncertainty hampers prudent decision making about where and when to invest most intensively in species conservation programmes. Demystifying ecosystem responses to biodiversity alterations may be best achieved through the study of the interactions allowing biotic communities to compensate internally for population changes in terms of contributing to ecosystem function, or their intrinsic functional redundancy. Because individual organisms are the biologically discrete working components of ecosystems and because environmental changes are perceived at the scale of the individual, a mechanistic understanding of functional redundancy will hinge upon understanding how individuals' behaviours influence population dynamics in the complex community setting. Here, I use analytical and graphical modelling to construct a conceptual framework for predicting the conditions under which varying degrees of interspecific functional redundancy can be found in dynamic ecosystems. The framework is founded on principles related to food web successional theory, which provides some evolutionary insights for mechanistically linking functional roles of discrete, interacting organisms with the dynamics of ecosystems because energy is the currency both for ecological fitness and for food web commerce. Net productivity is considered the most contextually relevant ecosystem process variable because of its socioeconomic significance and because it ultimately subsumes all biological processes and interactions. Redundancy relative to productivity is suggested to manifest most directly as compensatory niche shifts among adaptive foragers in exploitation ecosystems, facilitating coexistence and enhancing ecosystem recovery after disturbances which alter species' relative abundances, such as extinctions. The framework further explicates how resource scarcity and environmental stochasticity may constitute 'ecosystem legacies' influencing the emergence of redundancy by shaping the background conditions for foraging behaviour evolution and, consequently, the prevalence of compensatory interactions. Because it generates experimentally testable predictions for a priori hypothesis testing about when and where varying degrees of functional redundancy are likely to be found in food webs, the framework may be useful for advancing toward the reliable knowledge of biodiversity and ecosystem function relations necessary for prudent prioritization of conservation programmes. The theory presented here introduces explanation of how increasing diversity can have a negative influence on ecosystem sustainability by altering the environment for biotic interactions and thereby changing functional compensability among biota--under particular conditions.     

Ecological Aspects of Evolution of the Plague Microbe Yersinia pestis and the Genesis of Natural Foci

A new hypothesis of the origin of the plague microbe in the Mongolian bobak (Marmota sibirica Radde, 1862) populations in Central Asia during the Pleistocene is based on the ideas of its relative phylogenetic recency. The Late Pleistocene cooling, which induced a deep freezing of the grounds in southern Siberia, Mongolia, and Manchuria, is considered as an inducer of speciation. The main ecological factors of the plague microbe evolution include the species specific behavior of the Mongolian bobak as it prepared to hibernate related to its occurrence in arid petrophytic landscapes and the larval parasitism of the flea Oropsylla silantiewi Wagn., 1898 in winter. Genesis of the plague foci is divided into two periods: natural-historical and biosocial. During the first period, the primary natural foci in Eurasia were formed and, during the second period, synanthropic (rat) and secondary natural foci appeared with the participation of humans in Africa, The New World, and on some tropical islands.     

Task specialization in two social spiders,Stegodyphus sarasinorum (Eresidae) and Anelosimus eximius (Theridiidae)

Understanding the social organization of group‐living organisms is crucial for the comprehension of the underlying selective mechanisms involved in the evolution of cooperation. Division of labour and caste formation is restricted to eusocial organisms, but behavioural asymmetries and reproductive skew is common in other group‐living animals. Permanently, social spiders form highly related groups with reproductive skew and communal brood care. We investigated task differentiation in nonreproductive tasks in two permanently and independently derived social spider species asking the following questions: Do individual spiders vary consistently in their propensity to engage in prey attack? Are individual spiders' propensities to engage in web maintenance behaviour influenced by their previous engagement in prey attack? Interestingly, we found that both species showed some degree of task specialization, but in distinctly different ways: Stegodyphus sarasinorum showed behavioural asymmetries at the individual level, that is, individual spiders that had attacked prey once were more likely to attack prey again, independent of their body size or hunger level. In contrast, Anelosimus eximius showed no individual specialization, but showed differentiation according to instar, where adult and subadult females were more likely to engage in prey attack than were juveniles. We found no evidence for division of labour between prey attack and web maintenance. Different solutions to achieve task differentiation in prey attack for the two species studied here suggest an adaptive value of task specialization in foraging for social spiders.     

Bioassays with freshwater snails Biomphalaria sp.: from control of hosts in public health to alternative tools in ecotoxicology

Ecotoxicology is the science responsible for the study of the adverse effects of chemicals on ecosystems considering biotic and abiotic components. Several invertebrate groups have long been used to evaluate the aquatic toxicity of chemical compounds. Among these organisms, the microcrustaceans are the most recommended in Brazilian and international protocols (e.g. Daphnia sp. and Ceriodaphnia sp.). Until the beginning of the 1990s, the use of mollusks with ecotoxicological purposes was non-existent, except for the species tested as target of molluscicides in public health studies. Since the second half of this same decade the tests with mollusks have begun to be disseminated in several countries, valuing endemic species and especially the scarcity of test species in benthic habitats. In the early 2000s, with the disclosure of the harmful effects of pollutants with endocrine disrupting properties, gastropods have begun to be used not to evaluate lethal effects, but rather to observe physiological effects such as reproduction and embryonic development. Since then, assays with these approaches, especially with freshwater snails Lymnaea stagnalis and Biomphalaria sp., have been considered to be innovative and highly sensitive, often more than those achieved with traditional groups of test organisms in ecotoxicology (such as microcrustaceans and fishes).     

Are ecosystems hormetic?

The phenomenon of hormesis has been observed mainly for the response of individual organisms to stress. A reasonable line of inquiry might explore the possibility of observing hormesis at other levels of ecological organization. This initial examination focuses on ecosystem hormesis. Explorations of hormetic responses of ecosystems to stress cannot be made independently of a fundamental concept of ecosystem. The scale‐dependence of ecosystem dynamics also influences whether an ecological disturbance is in reality a stressor. Ecosystem hormesis might be claimed if one or more components of an ecosystem exhibit hormesis. By this definition, ecosystem hormesis would be a trivial extension of hormesis observed for individual organisms. A non‐trivial extension of ecosystem hormesis would include the observation that integrated (i.e., holistic) measures of ecosystem structure or function displayed an hormetic response to an ecological stressor. Several such examples of ecosystem structural and functional hormesis are presented.     

Natural selection and neutral evolutionary processes contribute to genetic divergence in leaf traits across a precipitation gradient in the tropical oak Quercus oleoides

The impacts of drought are expanding worldwide as a consequence of climate change. However, there is still little knowledge of how species respond to long‐term selection in seasonally dry ecosystems. In this study, we used Q_ST‐F_ST comparisons to investigate (i) the role of natural selection on population genetic differentiation for a set of functional traits related to drought resistance in the seasonally dry tropical oak Quercus oleoides and (ii) the influence of water availability at the site of population origin and in experimental treatments on patterns of trait divergence. We conducted a thorough phenotypic characterization of 1912 seedlings from ten populations growing in field and greenhouse common gardens under replicated watering treatments. We also genotyped 218 individuals from the same set of populations using eleven nuclear microsatellites. Q_ST distributions for leaf lamina area, specific leaf area, leaf thickness and stomatal pore index were higher than F_ST distribution. Results were consistent across growth environments. Genetic differentiation among populations for these functional traits was associated with the index of moisture at the origin of the populations. Together, our results suggest that drought is an important selective agent for Q. oleoides and that differences in length and severity of the dry season have driven the evolution of genetic differences in functional traits.     

Phenotypic plasticity to light intensity in Arabidopsis thaliana: invariance of reaction norms and phenotypic integration

Phenotypic plasticity (the pattern of response of organisms to changes in environmental conditions) and phenotypic integration (the pattern of character correlations) are important components of our understanding of the evolution of complex phenotypes. Most studies published so far in this area have been conducted within populations with the express aim of predicting future response to evolutionary forces. However, among-population differentiation for plasticity and trait correlations are important indicators of recent past events that have shaped the currently observable phenotypes. We investigated variation in the reaction norms of several traits in a large number of accessions of Arabidopsis thaliana exposed to different levels of light quantity as well as the environmental lability of the corresponding across-population character variance–covariance matrix. Our results show that there is an astounding degree of inter-population variation for character means and very little variation for plasticity, in agreement with the idea that A. thaliana is a light-specialist often occurring in open, disturbed habitats. However, this plant also shows patterns of plasticity that are predicted to be adaptive based on functional ecological considerations, such as an increase in either specific leaf area or leaf number (but not both) under low light. We also demonstrate that the set of character correlations in A. thaliana is extremely stable to changes in light availability, contrary to previous findings in the same species when different environmental factors were considered. Several processes that might have been responsible for the observed patterns are discussed as a prelude to follow-up research on these problems.     

Hybrid dynamics in a species group of swallowtail butterflies

Hybrid zones provide unique natural laboratories for studying mechanisms of evolution. But identification and classification of hybrid individuals (F1, F2, backcross, etc.) can be complicated by real population changes over time as well as by use of different marker types, both of which challenge documentation of hybrid dynamics. Here, we use multiple genetic markers (mitochondrial DNA, microsatellites and genomewide single nucleotide polymorphisms) to re‐examine population structure in a hybrid zone between two species of swallowtail butterflies in western Canada, Papilio machaon and P. zelicaon. Our aim was to test whether their hybrid dynamics remain the same as found 30 years ago using morphology and allozymes, and we compared different genetic data sets as well as alternative hybrid identification and classification methods. Overall, we found high differentiation between the two parental species, corroborating previous research from the 1980s. We identified fewer hybrid individuals in the main zone of hybridization in recent years, but this finding depended on the genetic markers considered. Comparison of methods with simulated data sets generated from our data showed that single nucleotide polymorphisms were more powerful than microsatellites for both hybrid identification and classification. Moreover, substantial variation among comparisons underlined the value of multiple markers and methods for documenting evolutionarily dynamic systems.     

Cells in the system of multicelular organism from positions of non-linear dynamics

The organism physiological systems forming a hierarchic network with mutual dependence and subordination can be considered as systems with non-linear dynamics including positive and negative feedbacks. In the course of evolution there occurred selection of robust, flexible, modular systems capable for adaptive self-organization by non-linear interaction of components, which leads to formation of the ordered in space and time robust and plastic organization of the whole. Cells of multicellular organisms are capable for coordinated “social” behavior with formation of ordered cell assemblies, which provides a possibility of morphological and functional variability correlating with manifestations of the large spectrum of adaptive reactions. The multicellular organism is the multilevel system with hierarchy of numerous subsystems capable for adaptive self-organization; disturbance of their homeostasis can lead to pathological changes. The healthy organism regulates homeostasis, self-renewal, differentiation, and apoptosis of cells serving its parts and construction blocks by preserving its integrity and controlling behavior of cells. The systemic approach taking into account biological regularities of the appearance and development of functions in evolution of multicellular organisms opens new possibilities for diagnostics and treatment of many diseases.     

The ‘Design’ of Mediterranean Landscapes: A Millennial Story of Humans and Ecological Systems during the Historic Period

What makes the structure and dynamics of coupled natural and human systems difficult to interpret in the Mediterranean is the extreme diversity in space and time of both environments and human societies. The succession of civilizations that waxed and waned in the Mediterranean Basin over several millennia has had great impacts on biota and ecosystems everywhere in the basin. A complex ‘coevolution’ has been claimed to shape the interactions between ecosystem components and human societies. Two opposing schools of thought traditionally have considered the consequences of human pressures on Mediterranean ecosystems. The ‘Ruined Landscape’ or ‘Lost Eden theory’ argues that human action resulted in a cumulative degradation and desertification of Mediterranean landscapes. The second school argues that humans actually contributed to keeping Mediterranean landscapes diverse since the last glacial episode. With this debate in mind, I show the following: (1) One cannot understand the components and dynamics of current biodiversity in the Mediterranean without taking into account the history of human-induced changes; (2) The various systems of land use and resource management that provided a framework for the blossoming of Mediterranean civilizations also had profound consequences on the distribution and dynamics of species, communities, and landscapes; (3) The processes of domestication of plant and animal species, which first occurred in the eastern Mediterranean area some 10,000 years ago, contributed to the increase of certain components of biodiversity at several spatial scales. Positive and negative feedback cycles between cultural practices and natural systems at the local and regional levels have kept ecosystems robust and resilient; (4) Assuming that human action can, to a certain extent, be considered a large-scale surrogate for natural sources of ecosystem disturbance, such patterns give support to the diversity-disturbance hypothesis—specifically, intermediate levels of disturbance have promoted biological diversity; (5) Intraspecific adaptive variation increased as a result of human-induced habitat changes over millennia, resulting in bursts of differentiation during the later Holocene of local ecotypes and gene pools of domesticated and wild plant and animal species, with region-specific characters fitting them to local climate and environmental conditions. High intraspecific adaptive variation also arose from earlier natural processes of the Pleistocene, mainly from a combination of periodic refugia formation and climate dynamics. During the Holocene, the main sources of disturbance came increasingly from humans, specifically from the coupled cultural and natural modifications of community and landscape structure. It is concluded that a high degree of resilience of Mediterranean ecosystems resulted in a dynamic coexistence of human and natural living systems, which in some cases provided stability, while fostering diversity and productivity (Blondel and Aronson, 1999). The word “design” used in the title and elsewhere in this paper metaphorically indicates that the long-lasting influence of human impacts resulted in an unintentional shaping of individual components of landscapes.     

The hologenome concept: we need to incorporate function

Are we in the midst of a paradigm change in biology and have animals and plants lost their individuality, i.e., are even so-called ‘typical’ organisms no longer organisms in their own right? Is the study of the holobiont—host plus its symbiotic microorganisms—no longer optional, but rather an obligatory path that must be taken for a comprehensive understanding of the ecology and evolution of the individual components that make up a holobiont? Or are associated microbes merely a component of their host’s environment, and the holobiont concept is just a beautiful idea that does not add much or anything to our understanding of evolution? This article explores different aspects of the concept of the holobiont. We focus on the aspect of functional integration, a central holobiont property, which is only rarely considered thoroughly. We conclude that the holobiont comes in degrees, i.e., we regard the property of being a holobiont as a continuous trait that we term holobiontness, and that holobiontness is differentiated in several dimensions. Although the holobiont represents yet another level of selection (different from classical individual or group selection because it acts on a system that is composed of multiple species), it depends on the grade of functional integration whether or not the holobiont concept helps to cast light on the various degrees of interactions between symbiotic partners.     

Salt to conserve: a review on the ecology and preservation of hypersaline ecosystems

When it comes to the investigation of key ecosystems in the world, we often omit salt from the ecological recipe. In fact, despite occupying almost half of the volume of inland waters and providing crucial services to humanity and nature, inland saline ecosystems are often overlooked in discussions regarding the preservation of global aquatic resources of our planet. As a result, our knowledge of the biological and geochemical dynamics shaping these environments remains incomplete and we are hesitant in framing effective protective strategies against the increasing natural and anthropogenic threats faced by such habitats. Hypersaline lakes, water bodies where the concentration of salt exceeds 35 g/l, occur mainly in arid and semiarid areas resulting from hydrological imbalances triggering the accumulation of salts over time. Often considered the ‘exotic siblings’ within the family of inland waters, these ecosystems host some of the most extremophile communities worldwide and provide essential habitats for waterbirds and many other organisms in already water-stressed regions. These systems are often highlighted as natural laboratories, ideal for addressing central ecological questions due to their relatively low complexity and simple food web structures. However, recent studies on the biogeochemical mechanisms framing hypersaline communities have challenged this archetype, arguing that newly discovered highly diverse communities are characterised by specific trophic interactions shaped by high levels of specialisation. The main goal of this review is to explore our current understanding of the ecological dynamics of hypersaline ecosystems by addressing four main research questions: (i) why are hypersaline lakes unique from a biological and geochemical perspective; (ii) which biota inhabit these ecosystems and how have they adapted to the high salt conditions; (iii) how do we protect biodiversity from increasing natural and anthropogenic threats; and (iv) which scientific tools will help us preserve hypersaline ecosystems in the future? First, we focus on the ecological characterisation of hypersaline ecosystems, illustrate hydrogeochemical dynamics regulating such environments, and outline key ecoregions supporting hypersaline systems across the globe. Second, we depict the diversity and functional aspects of key taxa found in hypersaline lakes, from microorganisms to plants, invertebrates, waterbirds and upper trophic levels. Next, we describe ecosystem services and discuss possible conservation guidelines. Finally, we outline how cutting-edge technologies can provide new insights into the study of hypersaline ecology. Overall, this review sheds further light onto these understudied ecosystems, largely unrecognised as important sources of unique biological and functional diversity. We provide perspectives for key future research avenues, and advocate that the conservation of hypersaline lakes should not be taken with ‘a grain of salt’.     

用功能生态学方法揭示峨眉髭蟾蝌蚪和棘腹蛙蟾蝌蚪共存的原因

采用功能生态学的研究方法,选取了3个指标(功能丰富度、功能定位和功能生态位的重叠程度)对峨眉山清音阁黑龙江中峨眉髭蟾蝌蚪和棘腹蛙蝌蚪共存的原因进行了解析。结果表明,峨眉髭蟾蝌蚪功能丰富度的实测值远远小于棘腹蛙蝌蚪。采用自助法随机抽样15和51个个体时,峨眉髭蟾蝌蚪的功能丰富度值显著小于棘腹蛙蝌蚪。同时,非参数多元方差分析的结果显示二者的功能定位存在显著差异。此外,二者功能生态位重叠程度的实测值与采用自助法计算所得值均极低。以上结果表明二者的功能生态位存在明显分化,表现为峨眉髭蟾蝌蚪个体较大,开口较小,眼间距较大,尾部发达且多采用背腹弯曲的游泳模式;而棘腹蛙蝌蚪开口较大,眼睛间距较小,尾部较为短小且多采用脊椎弯曲的游泳模式。鉴于功能性状可以反映生物体对生长环境的响应,推测两种蝌蚪主要栖息地的微生境也存在分化。