The influence of natural selection and sexual selection on the tails of sea-snakes (Laticauda colubrina)

Many phenotypic traits perform more than one function, and so can influence organismal fitness in more than one way. Sexually dimorphic traits offer an exceptional opportunity to clarify such complexity, especially if the trait involved is subject to natural as well as sexual selection, and if the sexes differ in ecology as well as reproductive behaviour. Relative tail length in sea-snakes fulfils these conditions. Our field studies on a Fijian population of yellow-lipped sea kraits ( Laticauda colubrina ) show that relative tail lengths in male sea kraits have strong consequences for individual fitness, both via natural and sexual selection. Males have much longer tails (relative to snout-vent length) than do females. Mark-recapture studies revealed a trade-off between growth and survival: males with relatively longer tails grew more slowly, but were more likely to survive, than were shorter-tailed males. A male snake's tail length relative to body length influenced not only his growth rate and probability of survival, but also his locomotor ability and mating success. Relative tail length in male sea kraits was thus under a complex combination of selective forces. These forces included directional natural selection (through effects on survival, growth and swimming speed) as well as stabilizing natural selection (males with average-length tails swam faster) and stabilizing sexual selection (males with average-length tails obtained more matings). In contrast, our study did not detect significant selection on relative tail length in females. This sex difference may reflect the fact that females use their tails primarily for swimming, whereas males also must frequently use the tail in terrestrial locomotion and in courtship as well as for swimming.     

Rupert Riedl and the re-synthesis of evolutionary and developmental biology: body plans and evolvability

This paper reviews the scientific career of Rupert Riedl and his contributions to evolutionary biology. Rupert Riedl, a native of Vienna, Austria, began his career as a marine biologist who made important contributions to the systematics and anatomy of major invertebrate groups, as well as to marine ecology. When he assumed a professorship at the University of North Carolina in 1968, the predominant thinking in evolutionary biology focused on population genetics, to the virtual exclusion of most of the rest of biology. In this atmosphere Riedl developed his "systems theory" of evolution, which emphasizes the role of functional and developmental integration in limiting and enabling adaptive evolution by natural selection. The main objective of this theory is to account for the observed patterns of morphological evolution, such as the conservation of body plans. In contrast to other "alternative" theories of evolution, Riedl never denied the importance of natural selection as the driving force of evolution, but thought it necessary to contextualize natural selection with the organismal boundary conditions of adaptation. In Riedl's view development is the most important factor besides natural selection in shaping the pattern and processes of morphological evolution.     

Australia's place in the global restoration challenge: Interview with Richard Hobbs

Summary   This interview with Professor Richard Hobbs, a prominent Australian researcher, professor and journal editor, traces his involvement in ecology and the relatively new disciplines of landscape ecology and restoration ecology. Born and educated as a plant ecologist in Scotland, Richard undertook postdoctoral research in the USA before taking up a series of research positions in Australia that steered him towards landscape ecology and restoration ecology. Having maintained an interest and involvement in international organizations, Richard provides comment in this interview on the progress of ecological restoration practice in Australasia compared to North America and comments on the need for ensuring research in these disciplines is strongly linked to management, is as broadly relevant as possible, and, is carried out at appropriate scales.     

Sources for Imanishi Kinji’s views of sociality and evolutionary outcomes

Prior to the contribution of genetics or the modern evolutionary synthesis (MES) to natural selection theory, social ecologists searched for factors in addition to natural selection that could influence species change. The idea that sociality, not just biology, was important in determining evolutionary outcomes was prevalent in research in social ecology in the 1920s and 1930s. The influence of ‘tradition’ (or the transmission of learned behaviours between generations) and the view that animals are active in selecting their own environments, rather than passive organisms acted upon by chance, were given as much attention as natural selection theory in European ecology, while animal aggregation and cooperation studies were pursued in America. Imanishi Kinji’s personal library and his scientific notes and papers reveal that he was well aware of this literature and had been profoundly influenced by these earlier viewpoints prior to writing his view of nature in his first book, Seibutsu no Sekai (The World of Living Things, 1941). Evidence is presented to show that he developed his theories based partly on early western debates in social ecology while finding inspiration and a way to express his views in the writings of philosopher Nishida Kitarō and, perhaps, General J C Smuts. One of Imanishi’s lasting contributions is in the demonstrated results of over 40 years of subsequent ecological and ethological research by Imanishi and those trained by him that maintained the broader viewpoints on evolution that had been dropped from the western corpus of research by the 1950s. The current attempt to again get beyond natural selection theory is reflected in debates surrounding genetic and cultural evolution of cooperation, the biology of ‘traditions’ and the idea of ‘culture’ in animal societies. Imanishi Kinji is the Japanese name order, with family name first. Other Japanese names in the text are also written with family name first. A modified version of this paper appeared in Japanese in Seibutsu Kagaku, Vol. 57 No. 3, April 2006, pp 142–149.     

Does Darwinism really contribute to ecology

The author questions Ghilarov's (2003) claim that Darwinism has high explanatory power in ecology. He is agree with S.V. Meyen who believed that beside synthetic theory of evolution (the popular variant on Darwinism) other explanations of evolution are possible. It is emphasized that several processes (e.g., diversification and unification of species at one trophic level, as well as individual and diffusive coadaptations of species of different levels) can contribute to community evolution. Communities cannot be considered as units of natural selection.     

Darwinism as a constraint of ecological pluralism

In his respond to critical remarks of Mirkin (2003), the author claims that pluralism in ecology is not only its strength but also a weakness. Contemporary ecology became less pluralistic and this can be considered as good sign of maturing science. Ecological pluralism can be exemplified by the coexistence in 1920-30s of two different approaches to plant community: that of Frederic Clements in USA and that of Josias Braun-Blanquet in France. However the way to progress in this branch of ecology was paved rather by heretical ideas of Henry Gleason in USA and Ramensky in Russia (both authors independently developed non-holistic view of community as an assemblage of individualistically distributed species) than by "peaceful" coexistence of well-established schools, representatives of which tried not to interfere into argumentation of each other. Notable success in ecology of last decades was connected with several new methodologies, e.g. macroecology that concerned large scale of space and time. However Darwinism in its attempt to explain the order of nature referring to its origin remains the most universal and fruitful methodology of ecology. The success of Darwinism in ecology is understandable because this generalizing theory is based on the same universal principles that underlie the survival of any population. Ecologists and evolutionary biologists trying to understand various natural patterns actually deal with the same fundamental laws, i.e. exponential population growth, limitation of this growth by resource shortage and/or press of predators, the existence of individual variability in survival, etc.     

The life and work of Hiroya Kawanabe: the priest ecologist

Hiroya Kawanabe was born the son of a Buddhist priest and teacher of Japanese literature, who died when Kawanabe was very young. Kawanabe also studied Buddhism by himself, and passed the examination to be a priest of his sect while still in high school. He studied zoology and ecology at Kyoto University and earned his doctorate under the guidance of Denzaburo Miyadi, a well-known Japanese ecologist, in 1960. During his academic career at Kyoto University, Kawanabe advanced to hold the chair of Animal Ecology as Professor in the Department of Zoology. Kawanabe's doctoral research concerned the social behavior and population ecology of the ayu, Plecoglossus altivelis, an amphidromous fish that lives in streams as adults and grazes algae. His research lead to the discovery that social structure changed from territoriality to schooling as population density increased, and also varied with changes in food and habitat. During this work, he pioneered the use of underwater observation to study ecolo gy of freshwater fishes in streams. Kawanabe also observed ayu social structure from the northern to southern limits of their range, and advanced the theory that the more stable territoriality in the Lake Biwa population was a relic social structure to guarantee food supply during earlier glacial periods when productivity was lower. Additional work on stream fishes in central Japan and Okinawa Island led Kawanabe to propose that interactions among individuals affect interspecific relationships, and thereby, community structure. Discussions with Charles Elton, the famous British ecologist of Oxford University, strengthened Kawanabe's view that communities could be best understood as the whole of interrelationships among organisms. Kawanabe advanced these ideas during a joint study he led with a host of Japanese and Zairean scientists on the fishes of Lake Tanganyika, beginning in 1979. This work, as well as additional research on Lake Biwa in Japan, led to a deeper understanding of the complexity of biotic interactions (including competition, predation, mutualism, commensalism, and indirect effects) that promote the high species diversity in these ecosystems. In addition to basic research, Kawanabe was part of research teams organized during the 1960s by D. Miyadi to study the effects of public works projects on natural environments and biota in Lake Naka-umi and Lake Biwa. During the late 1980s he expanded his network to an international venue, both by organizing and hosting important international ecological meetings in Japan, such as the Fifth International Ecological Congress, and by increasing his international activities to promote global biodiversity. In 1991, Kawanabe founded the Center for Ecological Research at Kyoto University to study the interrelationships among organisms and their environments. Recently retired from the University, he became Director General of the new Lake Biwa Museum in 1996, and continues to promote conservation of biodiversity worldwide through an international network of scientists and organizations.     

Darwin's "beloved barnacles": tough lessons in variation

In 1846, burdened by insecurity and self-doubt, and having been convinced that he needed to study some group of organisms closely, Darwin embarked on an eight-year odyssey in the protean and perplexing world of barnacles. At the time, he was searching for evidence in support of his theory of evolution by natural selection. In the course of his long study of barnacles, however, he was not just validating his preexisting theoretical system, but was also modifying his views on such fundamental aspects as the universality of individual variation, which is the focus of this paper. According to this notion, the members of any population of living things are expected to exhibit sufficient differences from one another for natural selection to operate. By emphasizing the theoretical value of the barnacle project, my analysis contributes to the historiographic tradition which highlights the significance of the period between the first comprehensive formulation of the theory of evolution by natural selection in 1844 and its urgent publication in the late 1850s. In the course of these years, Darwin's theory was not just accumulating empirical laurels, but was also expected to adapt to a changing conceptual landscape.     

作物生产是一个种群过程

从分析自然选择和人工选择在目标和方向上的重大差异入手,指出自然选择的核心是生存竞争,其结果产生个体优势;人工选择的方向是群体优势,其目标是获得较高的经济产量。作物是典型的人工选择的产物,研究作物的生产过程就庆当以降低个体的竞争能力和提高繁殖分配为主,据此,对小麦植株高度,根系特征以及种群生态学在农业中的应用进行了讨论     

Stage-structured cycles generate strong fitness-equalizing mechanisms

For many organisms, rates of reproduction, growth and mortality depend on the amount of resources that an individual consumes. When resource abundances fluctuate through space and time, the realized life-history of an individual can change dramatically depending on the dynamics experienced. Previous studies have investigated the influence of resource-dependent rates on population dynamics, but none have considered how the feedback between non-equilibrium resource dynamics and resource-dependent life-histories influence natural selection and the maintenance of genetic diversity within populations. Here we demonstrate that different patterns of resource dynamics have a strong impact on natural selection in organisms with resource-dependent life-histories. Small-amplitude consumer-resource cycles, lead to lower rates of natural selection than do large-amplitude consumer-resource cycles. Parameterizing the model for a Daphnia-algal system, we demonstrate that resource-dependent life-history can explain the recently published observation that selection among Daphnia genotypes changed depending on the pattern of algal resource fluctuations. The characteristically asexual reproduction of Daphnia allows us to draw a much-needed link to the large body of competition theory that has emerged from community ecology. Our results reveal that the common ecological features of resource-dependent life-history and ontogenetic size-structure generate strong fitness equalizing mechanisms that likely contribute to the maintenance of diversity in natural systems. Electronic Supplementary Material Supplementary material is available in the online version of this article at accessible for authorized users.     

Tree size distributions in an old-growth temperate forest

Despite the wide variation in the structural characteristics in natural forests, tree size distribution show fundamental similarities that suggest general underlying principles. The metabolic ecology theory predicts the number of individual scales as the −2 power of tree diameter. The demographic equilibrium theory predicts tree size distribution starting from the relationship of size distributions with growth and mortality at demographic equilibrium. Several analytic predictions for tree size distributions are derived from the demographic equilibrium theory, based on different growth and mortality functions. In addition, some purely phenomenological functions, such as polynomial function, have been used to describe the tree size distributions. In this paper, we use the metabolic ecology theory, the demographic equilibrium theory and the polynomial function to predict the tree size distribution for both the whole community and each species in an old-growth temperate forest in northeastern China. The results show that metabolic ecology theory predictions for the scaling of tree abundance with diameter were unequivocally rejected in the studied forest. Although these predictions of demographic theory are the best models for most of the species in the temperate forest, the best models for some species ( Tilia amurensis , Quercus mongolica and Fraxinus mandshurica ) are compound curves (i.e. rotated sigmoid curves), best predicted by the polynomial function. Hence, the size distributions of natural forests were unlikely to be invariant and the predictive ability of general models was limited. As a result, developing a more sophisticated theory to predict tree size distributions remains a complex, yet tantalizing, challenge.     

Multilevel selection 1: Quantitative genetics of inheritance and response to selection

Interaction among individuals is universal, both in animals and in plants, and substantially affects evolution of natural populations and responses to artificial selection in agriculture. Although quantitative genetics has successfully been applied to many traits, it does not provide a general theory accounting for interaction among individuals and selection acting on multiple levels. Consequently, current quantitative genetic theory fails to explain why some traits do not respond to selection among individuals, but respond greatly to selection among groups. Understanding the full impacts of heritable interactions on the outcomes of selection requires a quantitative genetic framework including all levels of selection and relatedness. Here we present such a framework and provide expressions for the response to selection. Results show that interaction among individuals may create substantial heritable variation, which is hidden to classical analyses. Selection acting on higher levels of organization captures this hidden variation and therefore always yields positive response, whereas individual selection may yield response in the opposite direction. Our work provides testable predictions of response to multilevel selection and reduces to classical theory in the absence of interaction. Statistical methodology provided elsewhere enables empirical application of our work to both natural and domestic populations.     

Ecological aspects of the evolutionary processes

Darwin in his On the Origin of species made it clear that evolutionary change depends on the combined action of two different causes, the first being the origin of genetically based phenotypic variation in the individual organisms comprising the population and the second being the action of selective agents of the external environment placing demands on the individual organisms. For over a century following Darwin, most evolutionists focused on the origin of inherited variation and its transmission; many workers continue to regard genetics to be the core of evolutionary theory. Far less attention has been given to the exact nature of the selective agents with most evolutionists still treating this cause imprecisely to the detriment of our understanding of both nomological and historical evolutionary theory. Darwin was vague in the meaning of his new concept of "Natural Selection," using it interchangeably as one of the causes for evolutionary change and as the final outcome (= evolutionary change). In 1930, natural selection was defined clearly as "non-random, differential reproduction of genes" by R. Fisher and J.B.S. Haldane which is a statement of the outcome of evolutionary process and which omits mention of the causes bringing about this change. Evolutionists quickly accepted this outcome definition of natural selection, and have used interchangeably selection both as a cause and as the result of evolutionary change, causing great confusion. Herein, the details will be discussed of how the external environment (i.e., the environment-phenotype interaction) serves as selective agents and exerts demands on the phenotypic organisms. Included are the concepts of fitness and of the components of fitness (= adaptations) which are respectively (a) survival, (b) direct reproductive and (c) indirect reproductive features. Finally, it will be argued that historical-narrative analyses of organisms, including classification and phylogenetic history, are possible only with a full understanding of nomological evolutionary theory and with functional/adaptive studies of the employed taxonomic features in addition to the standard comparative investigations.     

Synthesis and Selection: Wynne-Edwards' Challenge to David Lack

David Lack of Oxford Universityand V.C. Wynne-Edwards of Aberdeen Universitywere renowned ornithologists with contrastingviews of the modern synthesis which deeplyinfluenced their interpretation and explanationof bird behavior. In the 1950's and 60's Lackbecame the chief advocate of neo-Darwinism withrespect to avian ecology, while Wynne-Edwardsdeveloped his theory of group selection. Lack's position was consistent with thedeveloping focus on individual leveladaptation, which was a core concept of themodern synthesis. Alternatively, Wynne-Edwardsviewed the emphasis on populations as the mostimportant development provided by the modernsynthesis. In this paper, I present thedevelopment of these two positions and tracetheir roots in the literature of the synthesis.Through an analysis of Lack's 1966 critique ofWynne-Edwards I conclude that Wynne-Edwardswas, in many ways, justified in his pursuit ofgroup level explanations.     

Gulliver's further travels: the necessity and difficulty of a hierarchical theory of selection.

For principled and substantially philosophical reasons, based largely on his reform of natural history by inverting the Paleyan notion of overarching and purposeful beneficence in the construction of organisms, Darwin built his theory of selection at the single causal level of individual bodies engaged in unconscious (and metaphorical) struggle for their own reproductive success. But the central logic of the theory allows selection to work effectively on entities at several levels of a genealogical hierarchy, provided that they embody a set of requisite features for defining evolutionary individuality. Genes, cell lineages, demes, species, and clades-as well as Darwin''s favoured organisms-embody these requisite features in enough cases to form important levels of selection in the history of life. R. A. Fisher explicitly recognized the unassailable logic of species selection, but denied that thsi real process could be important in evolution because, compared with the production of new organisms within a species, the origin of new species is so rare, and the number of species within most clades so low. I review this and other classical arguments against higher-level selection, and conclude (in the first part of this paper) that they are invalid in practice for interdemic selection, and false in principle for species selection. Punctuated equilibrium defines the individuality of species and refutes Fisher''s classical argument based on cycle time. In the second part of the paper, I argue that we have failed to appreciate the range and power of selection at levels above and below the organismic because we falsely extrapolate the defining properties of organisms to these other levels (which are characterized by quite different distinctive features), and then regard the other levels as impotent because their effective individuals differ so much from organisms. We would better appreciate the power and generality of hierarchical models of selection if we grasped two key principles: first, that levels can interact in all modes (positively, negatively, and orthogonally), and not only in the negative style (with a higher level suppressing an opposing force of selection from the lower level) that, for heuristic and operational reasons, has received almost exclusive attention in the existing literature; and second, that each hierarchical level differs from all others in substantial and interesting ways, both in the style and frequency of patterns in change and causal modes.     

Unifying ecology and macroevolution with individual‐based theory

A contemporary goal in both ecology and evolutionary biology is to develop theory that transcends the boundary between the two disciplines, to understand phenomena that cannot be explained by either field in isolation. This is challenging because macroevolution typically uses lineage‐based models, whereas ecology often focuses on individual organisms. Here, we develop a new parsimonious individual‐based theory by adding mild selection to the neutral theory of biodiversity. We show that this model generates realistic phylogenies showing a slowdown in diversification and also improves on the ecological predictions of neutral theory by explaining the occurrence of very common species. Moreover, we find the distribution of individual fitness changes over time, with average fitness increasing at a pace that depends positively on community size. Consequently, large communities tend to produce fitter species than smaller communities. These findings have broad implications beyond biodiversity theory, potentially impacting, for example, invasion biology and paleontology.     

Sexual selection: Another Darwinian process

Why was sexual selection so important to Darwin? And why was it de-emphasized by almost all of Darwin's followers until the second half of the 20th century? These two questions shed light on the complexity of the scientific tradition named “Darwinism”. Darwin's interest in sexual selection was almost as old as his discovery of the principle of natural selection. From the beginning, sexual selection was just another “natural means of selection”, although different from standard “natural selection” in its mechanism. But it took Darwin 30 years to fully develop his theory, from the early notebooks to the 1871 book The Descent of Man, and Selection in Relation to Sex. Although there is a remarkable continuity in his basic ideas about sexual selection, he emphasized increasingly the idea that sexual selection could oppose the action of natural selection and be non adaptive. In time, he also gave more weight to mate choice (especially female choice), giving explicit arguments in favor of psychological notions such as “choice” and “aesthetic sense”. But he also argued that there was no strict demarcation line between natural and sexual selection, a major difficulty of the theory from the beginning. Female choice was the main reason why Alfred Russel Wallace, the co-discoverer of the principle of natural selection, engaged in a major controversy with Darwin about sexual selection. Wallace was suspicious about sexual selection in general, trying to minimize it by all sorts of arguments. And he denied entirely the existence of female choice, because he thought that it was both unnecessary and an anthropomorphic notion. This had something to do with his spiritualist convictions, but also with his conception of natural selection as a sufficient principle for the evolutionary explanation of all biological phenomena (except for the origin of mind). This is why Wallace proposed to redefine Darwinism in a way that excluded Darwin's principle of sexual selection. The main result of the Darwin–Wallace controversy was that most Darwinian biologists avoided the subject of sexual selection until at least the 1950 s, Ronald Fisher being a major exception. This controversy still deserves attention from modern evolutionary biologists, because the modern approach inherits from both Darwin and Wallace. The modern approach tends to present sexual selection as a special aspect of the theory of natural selection, although it also recognizes the big difficulties resulting from the inevitable interaction between these two natural processes of selection. And contra Wallace, it considers mate choice as a major process that deserves a proper evolutionary treatment. The paper's conclusion explains why sexual selection can be taken as a test case for a proper assessment of “Darwinism” as a scientific tradition. Darwin's and Wallace's attitudes towards sexual selection reveal two different interpretations of the principle of natural selection: Wallace's had an environmentalist conception of natural selection, whereas Darwin was primarily sensitive to the element of competition involved in the intimate mechanism of any natural process of selection. Sexual selection, which can lack adaptive significance, reveals this exemplarily.     

Styles of scientific reasoning: Adolf Remane (1898–1976) and the German evolutionary synthesis

Adolf Remane is widely considered to have been one of the most influential German zoologists of the 20th Century, yet Ernst Mayr persistently characterized him as an idealistic morphologist, that is, a typologist unable to understand population genetics or indeed Darwinian theory. This stands in sharp contrast to Mayr's praise for Bernhard Rensch as one of the most important German contributors to the Modern Synthesis of evolutionary theory. Remane's style of scientific reasoning is analysed in his writings on microsystematics, ecology, comparative morphology and phylogenetics and found to be highly consistent throughout these varied fields of research, while differing fundamentally from the eminently statistical foundations of both population genetics and natural selection theory that were embraced by Mayr. A comparative analysis of Rensch's understanding of science in general, and biology in particular, shows him to share core values with Remane, both authors rooted in the Mandarin tradition of the German professoriate. Biographical and socio‐political factors appear to have influenced Mayr's contrasting perception of Remane and Rensch, one that would influence later biologists and historians of science.     

Alternative allergy and the General Medical Council.

In July 1992 Dr Keith Mumby, a clinical ecologist, appeared before the professional conduct committee of the General Medical Council on five charges to do with his practice of clinical ecology. He was found guilty of two of the charges--touting for publicity and failing to give a patient adequate medical attention--and admonished. The GMC failed, however, to address the issue of the nature of Mumby''s treatments--clinical ecology itself. This is based on the idea that some patients are unusually susceptible to their environment, the diagnosis and treatment are based on an unstandardised provocation-neutralisation test. A variety of medical bodies have failed to find scientific foundation for the technique. The GMC''s policy on advertising services to patients is inconsistent, and in this case it has shown a regrettable reluctance to deal with the issue of treatments that are not scientifically validated.     

Environmental Security as a Case Study in Industrial Ecology

Environmental security is the integration of environmental and national security considerations at a national policy level. It is a relatively new and still somewhat contentious concept, although in some countries, such as the United States, it is increasingly being embedded in traditional security and foreign policy institutions. It is of interest to the industrial ecologist for several reasons. First, from the methodological perspective, environmental security issues are frequently complex, multidisciplinary, and multiscalar in both temporal and geographic dimensions. They are thus good opportunities to apply existing industrial ecology tools such as industrial metabolism stock and flow studies, as well as to support the development of new industrial ecology methods. Second, environmental security offers an important case study of an important fundamental industrial ecology dynamic: the movement of environment from overhead to strategic for society. This process occurs at many different scales, from implementation of design for environment methodologies within firms to integration of environmental and trade considerations in the World Trade Organization; and it is important for the industrial ecologist to begin to understand its underlying dynamics. Finally, national security is the quintessential raison d'etre of the national state. Accordingly, the integration of environmental considerations into national security policies and institutions, using industrial ecology methodologies and patterns of analysis, is a significant validation of the field.