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Biology and Philosophy 15: 211238, 2000. 2000 Kluwer Academic Publishers. Printed in the Netherlands.

Generalization in Ecology and Evolutionary Biology: From Hypothesis to Paradigm


KARI VEPSLINEN
Department of Ecology and Systematics P.O. Box 17 FIN-00014 University of Helsinki Finland

JOHN R. SPENCE
Department of Entomology University of Alberta Edmonton Canada

Abstract. We argue that broad, simple generalizations, not specically linked to contingencies, will rarely approach truth in ecology and evolutionary biology. This is because most interesting phenomena have multiple, interacting causes. Instead of looking for single universal theories to explain the great diversity of natural systems, we suggest that it would be protable to develop general explanatory frameworks. A framework should clearly specify focal levels. The process or pattern that we wish to study denes our level of focus. The set of potential and actual states at the focal level interacts with conditions at the contiguous lower and upper levels of organization, through sets of many-to-one and one-to-many connections. The number of initiating conditions and their permutations at the lower level dene the potential states at the focal level, whereas the actual state is constrained by the upper-level boundary conditions. The most useful generalizations are explanatory frameworks, which are road maps to solutions, rather than solutions themselves. Such frameworks outline what is understood about boundary conditions and initiating conditions so that an investigator can pick and choose what is required to effectively understand a specic event or situation. We discuss these relationships in terms of examples involving sex ratio and mating behavior, competitive hierarchies, insect life-histories and the evolution of sex. Key words: bottom-up pluralism, competitive hierarchies, contingency, explanation, generalization, life-history, many-to-one, mating behavior, multiple causation, nested hierarchy, one-to-many, scale, sex, sex ratio, triadic system, truth

We argue for a strategy that sees the unity of the general and the particular through the explanation of patterns of variation that are themselves higher-order generalities that in turn reveal patterns of variation. R. Levins and R. Lewontin (1985, p. 141)

212 In a thoughtful and provocative essay, Levins and Lewontin (1985) argue that investigations of the general and the specic are opposite sides of the same coin, and that neither approach to ecology is sufcient for progress. Although we agree wholeheartedly with this stance, the usefulness of some generalizations being offered currently in the literature of ecology and evolutionary biology can be questioned. This paper explores the process of generalization in evolutionary ecology with the intent of understanding why some generalizations are useful and stimulating, while others appear to be unsuccessful, ill-conceived or even counterproductive. Our premise is that useful generalizations can be characterized by their structure and their relationship to specic details of an ecological system, and that these characteristics may be revealed through analysis. We hold that generalizations are powerful research tools used at several levels to understand relationships between patterns and explanatory processes in nature. We begin our essay by considering the relationship between generalization and understanding because we believe that we can justify generalization in ecology mainly in the context of improving our understanding of the broad fabric of life. After a cursory review of basic strategies of modeling, we suggest that one basic approach in particular produces useful, biological generalization. We discuss several examples, drawn mainly from our own work, to illustrate the use of generalizations as research tools in ecology. Finally, we tackle the thorny problem of relating generalization to scale, and close with some guidelines for making generalizations in ecology. Generalization and understanding In its simplest sense, to generalize is to assert the existence of a pattern over an entire system, after observing only a subset of the system. This pattern, then, can be taken to unite the objects in the system in some dimension, although they may be clearly disparate in other ways. Effective generalizations establish some useful connection between objects of the system, and thus help us to reduce, often through abstraction, the complexity of the system of our attention and to see the main connections to the larger systems from which it is drawn. Ecologists commonly generalize about characteristics of levels in the biological hierarchy (e.g., populations, communities, ecosystems), or about elements that may be non-hierarchically organized within those levels (e.g., age distributions, food webs, nutrient cycles). We also seek to generalize about systems that have both ecological and historical dimensions (e.g., taxa, seres). Of course, for pragmatic reasons relating to the control of nature, humans desire ecological generalizations that are simultaneously simple (i.e., easy to understand or inexpensive to implement) and powerful (i.e., explain

213 all cases or always work). However, given the complexity of ecological systems, such generalizations are hard to come by. For most of us, understanding conveys some feeling that our immediate observations or thoughts can be connected to some larger system the elements of which we cannot experience both simultaneously and in detail while making observations in nature. We understand by explaining these connections in periods during which we analyze sensory information. It is crucial that we know enough about the larger system to see how our notions can be settled within it. The scale of generalization required relates in some sense to the size of our notions. For many ecologists, one community, one taxon or even one population is the relevant frame of reference, whereas for others larger-scale generalizations are necessary. Given the hierarchical structure of biological information mentioned above, it is clear that some notions are most appropriately connected to lower-scale concepts. The levels at which ecologists work are largely a matter of personal choice and ability, although there is prejudice against work that is not connected to simple, broad-scale generalizations, and as a result such connections may be commonly forced. Generalizations vary in their level of abstraction and in the extent that they are directly connected to specic objects in the real world. At the most basic level, generalizations are the testable hypotheses that ecologists use to investigate particular processes; these usually involve minimal abstraction and provide a one-to-one match between parameters and phenomena that can be measured in nature. The less knowledge we have about a phenomenon of interest, the more such generalization helps in gaining a working knowledge of nature, but at the same time, the more likely the generalization is to fail. As Popper (1965) argues, bold conjecture puts our hypotheses to strong test. If they pass such tests, the ideas become more generally useful; if not, the ideas should be discarded or reworked. However, useful generalizations may be more abstract than strict hypotheses and propose relationships among classes of parameters that are not explicitly connected to specic systems. These generalizations are not at all testable in any particular system because part of the challenge is to connect the abstraction to measurable elements in each system. The concept of natural selection, for example, is an abstract notion that both Darwin and Wallace arrived at by seeing common threads in many systems, and then suggesting that such threads could be connected in a way to explain much of the diversity of life on earth. Although some (e.g., Peters 1976) point out that the concept of natural selection is not testable by particular experiment, and suggest that this shortcoming disqualies it as a scientic theory, such highly abstract generalizations are essential frameworks for asking more specic questions about nature. In principle, such frameworks are tested each time they are used

214 to explain a particular set of observations and they are discarded if another synthesis supporting a broader range of useful explanations becomes available. However, the ability of these frameworks to generate good questions is more central to their validity as scientic constructs than is the criterion of falsiability whether or not we choose to allow them to be called theories. For Kuhn (1962), expressions at relatively high levels of generalization in science are paradigms and these are essential for scientic progress. We hold that generalization in science, and in ecology in particular, includes the broad continuum between hypothesis and paradigm, and that the value of any generalization springs mainly from its usefulness in promoting research that leads to understanding. Thus, an insistence that generalizations be true seems to depend on the level at which they are offered. Low-level generalizations about particular systems, or hypotheses, must be immediately falsiable by experiment; they are formulated easily and we scurry to test them. Paradigms, on the other hand, are upper-level generalizations that promote the construction of hypotheses but that are not falsiable by experiment alone. When classes of similar things, in this case generalizations, are dened, it usually turns out that there is a continuum that connects them. Thus, there are useful generalizations, not strictly testable, that are broader than hypotheses but more specic than paradigms. The role of these generalizations, which we refer to as explanatory frameworks, is commonly misunderstood by ecologists. Perhaps the words hypothesis and theory have been worshipped too much by scientists in general. A main point of this essay is that these are only generalizations about nature that lie somewhere along a continuum of abstraction. When this is recognized, the common rejection of a new generalization because it is not testable in the narrow sense seems absurd. Some ideas must be worked with and developed before they t into the hypothetico-deductive context of contemporary science. Their validity as scientic constructs springs entirely from the interest that they generate among scientists. We fear that science can be trivialized by overriding emphasis on the narrow interaction between hypothesis and test. Somewhat paradoxically, both the need to generalize and the aws in biological generalizations spring from the same basic fact the staggering diversity of life. Not only are ecological systems potentially huge, but their possible classications are complex. This means that effective biological generalizations are bounded, and we can expect them to fail when invoked outside of those boundaries (Levins and Lewontin 1985). The dilemma for all scientists is that detailed knowledge supporting strong inference about particular systems is limited to bounded systems. For example, chlorouorocarbons (CFCs) were shown to be chemically inert and without biological

215 consequences over a fairly large spatio-temporal scale before their wide application. However, now that we have become painfully aware of effects of CFCs outside of the original boundaries, the original generalization about their safety seems virtually useless, and certainly is no longer invoked to support their continued use. Because the very structure of biological information is hierarchical, we necessarily lose track of one kind of information as we consider increasingly larger sets; however, we sometimes forget that we gain other information. A conceptual problem for ecologists is that this additional information may be several levels removed from the traditional biology which was the basis of their formal education. Such information, as for example woven together in the Gaia Hypothesis (Lovelock 1979, 1991), may have profound implications for our understanding of more local ecological processes (Lenton 1998). Important advances in ecology can be outside of our experience and in the words of Bonner (1969), they can be on a scale too large, so that, if anything, our experience becomes an impediment to our easy understanding, and we must learn we can no longer trust our senses with the commonplace information they provide. Of course, the trick is to connect events on the large scale to measurable predictions at scales at which we can work. Perhaps the most signicant advance in ecology during the past two decades is an increasing acceptance that multiple causation is the norm in ecological systems, and that therefore efforts to identify single factors to explain ecological processes in all systems are doomed to fail (Hairston 1989). For example, simple generalizations such as populations are regulated by specic key factors that operate with strong density-dependence, which were accepted by many in the 1960s, are now known to be bounded by both space and time. The apparent causal phenomena that we carefully measure in todays ecological play, may disappear from the stage in the next valley or in the next season. However, the general statements of yesterday may become tools in a search for unifying phenomena on other planes. Multiple causation is an obvious consequence of the many to one connections that result from the hierarchical structure of biological systems (Levins and Lewontin 1985). Research strategies The relationship between generalization and understanding may go seriously wrong in at least three basic ways. First, generalizations may y in the face of numerous facts. This error may result when authors are not sufciently aware of the literature about their problem or about the natural history of the group over which they generalize. This sort of error is usually detected quickly by specialists, although most taxon specialists can point to many

216 such errors perpetuated in textbooks about his particular group. Although there is tremendous pressure on ecologists to work on general propositions, we cannot make things general through assertion even if such assertion is published in prestigious journals. Second, generalizations fail when the information brought from one level cannot be matched to the terms in the generalization. On one hand, when rich, lower-scale information is related to generalizations insufciently detailed to allow direct connection of all relevant information brought forward from below, we feel lost, unable to nd landmarks within a fog of strange, abstract jargon. On the other hand, when too much irrelevant detail is carried up to the next level for generalization, we feel overwhelmed, unable to t all the information at hand into the classication provided. Without intervening levels of abstraction or work to reduce the detail of our data, connections to the generalization are fuzzy and little effective understanding is achieved. For example, take the statement heterogeneous environments are characterized by high rates of dispersal. However detailed and/or large our data set on dispersal would be, the link between the data and the generalization would evade our understanding simply because heterogeneity as a concept is too heterogeneous itself. Different kinds of environmental heterogeneity favor different kinds of life histories. Third, when generalizations are not effectively bounded, those who work on systems outside of the universe that is properly referenced do not nd the general concept useful. This is the source of feelings that many socalled general principles in ecology and evolutionary biology apply only to birds, butteries or owering plants. Biological generalizations may reect a tyranny of the most vocal or most abundant biologists, rather than reecting important features about the majority of taxa or ecological systems to which they may properly refer. A generalization is not proven useless simply because we have not understood or properly articulated its original domain. For example, consider the statement, seasonality selects for migration. The test data must be bound by phylogeny and functional morphology. Birds may escape wide-spread, large-scale vicissitudes by covering large stretches, and returning to their breeding grounds for the more benign season. However, because insects are less likely to survive through such journeys and time spans, their inclusion in the test data would seriously weaken the test of an otherwise well-dened generalization, although a few insect species do indeed undertake large-scale migrations. Another sort of error is encountered in statistical generalizations, that have become increasingly popular among empirically oriented ecologists. Where we assert, for example, that the usual behavior in taxon 1 is x as a shorthand version of saying that 60% of the species so far examined behave

217 that way, we provide a simple generalization. Although this may be used as a trivial sort of hypothesis by those working on taxon 1, the generalization is demonstrably wrong in 40% of the subsystem where the usual behavior is y. Where such errors are important, as in making management decisions about a specic member of taxon 1 or in forging a new generalization about the next higher taxon, little useful purpose has been served. Statistical generalizations can even be true, but still rejected as unworthy of attention by the scientic community at large. For example, rational zoologists reject, in the context of curriculum development, the usefulness of the generalization that the vast majority of known animals have six or eight legs, even though it is technically correct. In the broadest sense, we generalize by building mental models about nature that attempt to explain an observed pattern or process by simplifying the complexity of the real world. We feel comfortable with our explanations when the model mimics the pattern or process under investigation, even though this comfort may be unwarranted (Weizenbaum 1976). The rst step is to abstract the essential features of the system under investigation. Aspects of the system, thought to be non-essential for the purpose of the study, are excluded from the model by bounding it. Thus the process of modeling sacrices some information known about the system to increase understanding of particular aspects about which we seek to generalize. Levins (1966) visualized the tradeoffs in relation to three main approaches to modeling, which give differing emphasis to a models generality, realism and precision. Applied ecologists tend to sacrice generality to realism and precision. For example, sheries biologists have modeled population dynamics through statistical descriptions of population changes over many years. These descriptions lead to a multiparameter regression equation of best t to the real data. The regression model is used to predict future population changes. However, faithful description of the data does not qualify as an explanation for the pattern. True understanding of the dynamics may be achieved only by connecting them to some upper level generalizations about the factors that bring about changes in sh stocks. Although some broad correlations with todays decreasing catches are evident, few would suggest that we understand the dynamics of even the most commercially important sh populations. The arm-chair approach to ecology, often produces models sacricing realism to generality and precision. Assumptions of the models may be met by very few, particular biological systems. Mathematical simplicity and elegance are the main criteria for success. Abundant generalizations emerge, but these are only connectable to nature where the assumptions necessary for the mathematics are met. The approach has not attracted many good natural-

218 ists to theoretical ecology although it invites some theoreticians into ecology and later to become pretty fair naturalists. We believe that the third strategy, sacricing precision to obtain generality and realism, leads to the most effective generalizations. In this approach, directions of changes or differences between objects or systems are more important than exact magnitudes (Levins 1966, 1968, 1974; Puccia and Levins 1986). The loss of precise information about exact rates or numbers is compatible with the hierarchical structure of biological systems. Some examples of generalization We now turn to some examples of generalization, spanning the range from specic hypothesis to framework. Although we present mainly entomological examples drawn from our own areas of research, we believe that they support a broader analysis of generalization in ecology and evolutionary biology. We aim to illustrate how generalization allows effective coupling of focal-level results with the main threads of information brought up from lower levels and with an understanding of boundary conditions. We also show how they fail when such coupling is not achieved. 1. Mating behavior: Intersexual conict or not? Sperm is metabolically inexpensive to produce in large quantities, whereas production of large eggs is more costly. Based on this simple observation, behavioral ecologists generalize that males behave to maximize quantity of matings, and that females are much more choosy about partners in an attempt to maximize quality of their offspring. This generalization, springing from knowledge of celllevel, initiating conditions, has promoted a urry of research activity about the expected conict between males and females for mating opportunities. Even though signicant exceptions to the predictions have been found (e.g., paternal investment, sperm displacement; see Thornhill and Alcock (1983) for examples dealing with insects) most investigators still begin with the expectation that males will insist on mating repeatedly, whereas females will often resist. When the behavioral expectations are not met, we begin the search for upper-level constraints that may affect our expectations. Ecologists generally look to environmental heterogeneity to provide explanations for unexpected variation. In the case of mating behavior and most other problems, a simple appeal to global heterogeneity neither explains much, nor does it help us derive testable hypotheses applicable to any specic situation. Thus, we attempt to partition heterogeneity across factors that promote it such as predation, food availability, oviposition sites, operational sex ratio, etc. By invoking boundary conditions on the focal-level

219 processes of our interests, we are forced to specify the original generalization, i.e., to make it less general. As an example of this approach in the study of mating behavior, we discuss studies of variation of sex ratio in waterstriders (Gerridae). Males of many gerrid species succeed in getting females to mate repeatedly, despite the fact that 23 matings is sufcient for a female to fertilize her entire complement of eggs. We hypothesize that the basic intersexual conict, predicted by the very general theory above, will be less observable when the ratio of bachelor males to non-virgin females is high because, when all costs are considered, pressure on males to mate will be stronger than pressure on females to resist. The freedom of males to hunt for new mates is constrained by male-male competition for females, and under male-biased sex ratios, time in copula and that spent in post-copulatory guarding tend to be extended. The sperm of a male that does not guard may be displaced by sperm of a subsequent male who mates before the female oviposits (Arnqvist 1988; Rubenstein 1989). For females in male-biased situations, costs of strong harassment may outweigh costs of accepting an additional mating. It follows that females should resist less, and both females and males should agree in prolonging mating. As shown by several investigators (e.g., Arnqvist 1992a, b; Rowe 1992; Jablonski and Vepslinen 1995), such general expectations are met in experiments in Gerris species, when individuals in male-biased experiments are compared with individuals in even or female-biased ratios. It would be now tempting to generalize that variation of operational sex ratios (OSR) during mating explains mating behavior. But the generalization applies to an experimental environment where only mating-environment sex ratio was varied and all other potentially important boundary conditions were neglected. In the real world of waterstriders, relevant boundary conditions certainly include other factors such as risk of predation, and variation in food and microhabitat structure (Sih 1988; Sih et al. 1990; Rowe 1992), and patterns seen in nature may frequently differ from that suggested by simple generalizations about the effects of OSR. As an example of boundary conditions traditionally neglected in behavioral studies, we take the OSR experienced by males and females before the mating attempts by the male. It is known that the OSR varies much in time and space for waterstriders within a single pond. Thus an amorous male bumping to a female here and now may have a different OSR experience from that of the actual mating environment; the same is true for the female. When this new dimension in the boundary conditions is experimentally manipulated and controlled for, the effect of recent OSR experience of males and females can be shown to be more important than the OSR of the mating environment in affecting the length of mating, the proportion of postcopulatory guarding, and

220 the success rate of the males (Vepslinen and Savolainen 1995). Therefore, when we bring rich information in from nature, we may expect that it will not match the laboratory results. In a sense, the generalization fails for the second reason given above; we have pushed it beyond the original boundary conditions implicitly specied by the highly simplied laboratory environment. We do not achieve generality by looking, no matter how carefully and repeatedly, at one thread in a multicausational connection. As evident from above, it is not important whether the original generalization about behavior, deduced from a comparison of the metabolic cost of sperm and eggs, is right or wrong. It is a useful generalization, a good research tool, if it inspires interest in a problem and leads us eventually to a more satisfying understanding of nature; however, widely accepted generalizations may also retard development in the eld because of conservative social pressures. Part of this process of generalization is the generation of testable hypotheses of biological relevance. Results of experiments may be used to modify the generalization, often by stepping down to a lower level of generality, as in this example, to the population level where decisive variation for the factors affecting mating occurs. Through attention to the boundary conditions surrounding our explanatory framework, we may produce a set of new predictions, each capable of generating testable hypotheses. Complexity will grow at the expense of simple generality, but so will our connection with the reality of nature. To generalize effectively about ecological phenomena, given all the glorious complexity of a natural system, necessarily implies specication. Working out contingencies is essential.1 2. From individual behavior to structuring of communities. Many theories are connected only loosely to the natural world that can be perceived directly by human observers. We maintain that in order to build frameworks to help explain the operation of nature, we must become intimately familiar with our study objects on a variety of scales. Collecting, observing and reading widely about the groups that we study allow us to master the basic natural history that is necessary to reasonably bound generalizations. Admittedly this naturalizing stands outside of the scope usually embraced as science, but we hold that its recent de-emphasis in biological training is associated with proliferation of futile generalization in ecology. While striving towards understanding ecological processes, we use our data for existential statements and generalizations, which make the point that dening ecological phenomena is not trivial and deserves attention observations should be generalized to a level that becomes theoretically relevant (Haila 1988). In this vein, Vepslinen and Pisarski (1982) suggested on the basis of their long-time myrmecological experience that ant species can be ordered along a linear competitive

221 hierarchy, depending on their tendencies to indulge in erce frontal attacks in conict situations. The hierarchy consists of the following three basic levels: (1) territorial species which attack individuals of other colonies and species on their food territory; (2) encounter species which are not territorial, but defend concentrated food sources (e.g., aphid colonies); and (3) submissive species which defend only their nests but otherwise avoid aggressive encounters through evasive behavior. This simple conceptual framework allows predictions about which species combinations are probable in local ant assemblages. Territorial species should, by denition, exclude each other in space. Encounter species tend to come into conict with and be excluded by territorial species. However, submissive species may coexist with other ant species, given they are able to accommodate ecologically or behaviorally to the aggressive behavior of competitively superior species. In this work, the structure of ant communities is the focal level of interest. However, because the initiating conditions are at the level of individual behavior, experiments on the outcome of behavioral interactions among individuals of different ant colonies and species are necessarily done on a small scale, e.g., by observing behavior at baits. Findings that agree with the predictions of the competition hierarchy can be generalized to the level of communities. This produces predictions about the distribution pattern of ant species on a larger spatial scale than that of the bait experiments done to study the initiating conditions. Data can then be collected to test predictions about patterns of nest dispersion. The predictions generated by the above general framework can be tested with detailed analytical descriptions (Haila 1988) of the relations among the species of local assemblages. Here simple eld experiments within the tradition of hypothetico-deductive testing were used to test predictions about the frequency and outcome of aggressive encounters and to support the generalization on a small spatial and temporal scale (Savolainen and Vepslinen 1988, 1989; Savolainen 1990, 1991; Vepslinen and Savolainen 1990). After experimental support was obtained for the small-scale generalization, Savolainen et al. (1989) expanded the generalization to a larger scale on the basis of information in the literature. They suggested that multinest societies of wood ants (Formica rufa group), the species highest in the competition hierarchy, organize ant assemblages and, more generally, whole communities of predatory forest-oor invertebrates throughout the Holarctic taiga region. Unpublished evidence gathered so far about importance of wood ants in organizing communities, tends to discourage extension of predictions to the entire Holarctic. For unknown reasons, Nearctic wood ants lack the organizing power seen generally in the Palearctic. Perhaps for historical

222 reasons Nearctic wood ants have not reached the level of social organization required to have strong impact, or the reasons may well be at least partially ecological, depending on the exact mix of species. Nonetheless, the original theoretical framework still allows us to look for detailed reasons for the failure of the generalization, and to make theoretically justied comparisons (Haila 1988) between the taiga regions of the two continents. Such comparisons are needed to nd the right levels where the initiating and/or boundary conditions may differ between the two continents. 3. The habitat templet framework. Since the pioneering work of Lamont Cole (1954), attempts to understand the evolution of life-history traits have been a mainstream of ecology. MacArthur and Wilson (1967) found a common thread for (i.e., generalized) the process of life-history evolution by suggesting that many life-history traits evolved under selection pressures that could be described on a continuum from r- to K-selection, associated respectively with growing and stable populations. They developed these ideas primarily in order to explain differences in life-history characteristics of species on islands vs mainlands. However, the apparent power of the rK framework and its direct connection to other explanatory frameworks in population biology set the stage for generalization in two directions. First, there was urry of papers that further generalized application of the framework (e.g., Pianka 1970, 1972) by exploring its broader correlates and internal consistency, outside of its original domain (uncrowded, no-competition vs crowded, strong-competition environments; rigorously seasonal vs unseasonal climates; island-mainland comparisons). Second, a host of authors applied the generalization to interpret life-history patterns observed in particular taxa (e.g., McNaughton 1975; Satchell 1980; Spence and Scudder 1980; Vepslinen and Patama 1983). Although the generalization was useful for some specic systems, it was found wanting as an explanatory framework in others. In a landmark paper, Stearns (1976) reviewed life history theory and concluded that the concept of bet-hedging was a necessary addition to the r-K framework used for constructing hypotheses about life-history theory. Extension in this direction allowed predictions about life-history traits in situations where mortality and fecundity schedules uctuate. This important synthetic work broadened the domain of life-history theory, especially for vertebrates, and has been particularly successful in framing studies of the age distribution of reproductive effort. The important point, in the present context, is that the world is not, by design, either an r-K or bet-hedging universe, but these concepts are human-derived tools that express and facilitate our efforts to understand a huge and diverse array of natural history information

223 about life-history traits of animals. In a sense, these tools simply provide a bridge connecting life-history work to the theory of natural selection by proposing denite relationships among a reasonable number of parameters that can be identied in systems that exist in nature. Their utility springs from the strength of connection to the general theory on one end, and to a signicant number of specic biological systems on the other. Insects comprise the vast majority of described species, and yet the general framework for asking life-history questions brought together by Stearns (1976) is not particularly relevant for explaining some striking aspects of life-history variation among insects. These aspects are mainly concerned with how persistent insect species adjust and time reproduction to deal with risk encountered on scales sometimes different than those relevant for larger animals. The habitat templet approach as developed by Southwood (1977, 1988) and Greenslade (1983) includes both adult movement associated with dispersion of offspring, and diapause as components of life-history that evolve along with reproductive traits, thus focusing attention on adaptive tradeoffs not normally considered in work with vertebrates. The approach holds that life-histories evolve in an ecological space dened by three axes depicting variation in habitat characteristics: (i) a disturbance axis related to the r-K model, (ii) an adversity or productivity axis, and (iii) an axis related to biotic interactions. Southwoods (1977) stated aim is to provide a classication of life-histories with respect to habitat features to serve as a sort of periodic table to predict the traits of unstudied species that could be classied with respect to the habitats that they use. We emphasize that Southwoods habitat templet model is not a testable hypothesis that can be falsied with study of any particular system. Its validity as a scientic construct ows from its utility as a broad generalization made through abstraction about the environmental features important for evolution of life-history traits, particularly in insects. Such generalizations are frameworks for asking questions that allow us to understand more about particular natural systems. The habitat templet framework has contributed signicantly to the understanding of insect life-history evolution. It has been particularly relevant to our own work with semi-aquatic bugs, a group that shows an uncommonly high degree of life-history variation in movement and diapause traits that has traditionally been related to habitat features (summarized in Vepslinen 1978; Spence and Andersen 1994). Most temperate-zone gerrids use vegetated zones of shallow waterbodies as habitats (Andersen 1982). Because these may dry out or change dramatically in character during a single season, initial adaptive explanations for variation in movement and diapause traits were explained as correlations with habitat permanence, one axis in the templet model related to the r-K continuum (e.g., Vepslinen 1974, 1978;

224 Spence and Scudder 1980). These explanations, although fairly robust at the level scrutinized by non-specialists, left out signicant aspects of gerrid lifehistory variation, and this deciency was the subject of enthusiastic debate in many subsequent gatherings of waterstrider experts. The question was whether we could provide a better synthesis and formulate it in a way that could be linked to broader generalizations about life-history evolution. The answer is not yet clear but work is proceeding in several directions. Spence (1989) chose a path suggested by the habitat templet framework and tried to incorporate information about productivity and biotic interactions into our understanding. Work completed to date suggests that both food availability (Spence 1989 and in prep.) and the presence of natural enemies (Spence 1986a, b; Nummelin et al. 1988) can inuence the tness of gerrids employing different life-history tactics. We are now trying to establish whether patterns of variation in these general parameters are stable enough to provide consistent selection toward life-history correlates. In addition, Klingenberg and Spence (1997) showed that the two central trade-offs of modern life-history theory (Roff 1992; Stearns 1992) simply do not hold for at least one well-studied and common gerrid species. Specically, juvenile development time and adult body size are inversely proportional and larger females do not, on average, lay more eggs in G. buenoi. The best presently available hypothesis for this is that large body size is driven by strong selection for ight ability in this species which is characterized by use of unstable and unpredictable habitats. The so-called general theory fails, in this case, because its generalized underpinnings do not hold; these insects differ from other creatures in relation to life-history features captured by Southwoods (1977) generalizations about the habitat templet. Thus, it appears that general life-history has a domain that does not include all species, a fact noted by an early reviewer of the Klingenberg and Spence (1997) work who wrote, general life-history theory was not developed to deal with gerrids so what? Highly dispersive species, such as G. thoracicus, provide another challenge for models of life-history evolution. Here, it appears that differences in the mortality schedules of juveniles and adults have to be considered as suggested in the bet hedging model (Vepslinen 1978 and unpublished). In a highly variable and unpredictable set of environments, supercially described as r-selective, Finnish populations of G. thoracicus show high variance in juvenile mortality coupled with much lower and less variable mortality of reproducing adults. Adults of this species are long-lived, have lengthy pre-reproductive periods (Vepslinen and Patama 1983), and spread reproductive effort out in time and across habitats. Consequently the life history of G. thoracicus looks supercially closer to that of a traditional

225 K-selected organism than to that of an r-selected species, although the species seems to be a relatively poor competitor. The adaptive signicance of diapause and migration in this species can be understood only in relation to how the habitat templet inuences the relative uctuations of adult and juvenile mortality schedules. The ultimate outcomes of our research is not especially relevant for the present discussion. What is important is that the work itself has been stimulated by upper-level generalizations about the organization of relevant selective forces. For example, Southwoods synthetic generalization has supported formulation of testable hypotheses about particular systems of semi-aquatic bugs, and the results of experiments done to test these hypotheses promise to further hone the generalization by revealing the pattern of variation that surrounds it. The framework may not be particularly useful for study of relatively long-lived vertebrate species and other organisms for which ight in association with diapause are not a major adaptive response to seasonal habitat changes. It focuses attention on what are null adaptive sets in some species. However, through discovery and testing of satisfying explanations for life-history adaptations in a variety of creatures we may hope to simultaneously appreciate the diverse fabric of nature and discover effective generalizations on a higher level. Perhaps waterstriders arent so very different from birds, butteries and owering plants afterall. The generalization of a tapestry involves combining an abstract idea or general plan with the weaving of many diverse threads. 4. Apomictic vs sexual reproduction. In Masterpiece of Nature, Graham Bell (1982) reviews the paradox of sexuality, seeking to nd a universal theory that explains different modes of reproduction. He lists aptness, generality and simplicity as three aesthetic and even poetic criteria for judging rival explanations, and argues further that decisions among apt theories should be made on the grounds of generality and simplicity. Generality is held to be more important because it allows theory to enjoy wider application. In attempting to explain the presence of sexual or asexual reproduction among metazoans, Bell bases reasons from an axiom of perfection which contends that adaptation is precise, and that selection alone is a sufcient explanation of organic diversity. Essentially, he claims that by invoking accidents and history in our explanations, we resort to something akin to mythology. Bell ultimately gives way to provide a magnicent and more pluralistic summary about the probable evolution of sexuality which includes historical elements. However, we use his opening thesis and basic approach (as outlined above) to illuminate several difculties inherent in strictly deterministic, gene-level, selectionist approaches to constructing a single,

226 universal theory to explain sexuality. Our goal is to underscore approaches required for effective generalization. Although sexual reproduction seems to be an ancient character of metazoans, many species in higher taxa have returned to asexual reproduction, or deviated in some other way from amphimixis, or normal sexual reproduction. For example, apomictic lineages have dispensed with meiosis and syngamy so that reproduction is essentially cloning with progeny more or less genetically identical to parents and so that the level of heterozygosity is conserved. Cyclic parthenogenesis is characterized by alternation of phases of amphimixis and apomixis so that high diversity and heterozygosity are conserved or restored. In amphimixis, there is a potential for great variation in the degrees of heterozygosity and diversity, depending on population structure and the details of mating system. However, one feature standing clearly in contrast to apomictic reproduction, is that a parental genome is never transmitted intact to the offspring. Whether this bestows higher or lower tness to amphimictic organisms, relative to apomictic ones, must depend on the pattern of the environment relative to the constraints set by the cytology of each reproductive strategy. This reects an important problem for science. Although the ideal of modern science is universal explanation, it is problematic to deduce specics from generalities when the form of specics is profoundly inuenced by unique, local situations (Haila and Levins 1992). This logical problem compels evolutionary biologists to begin with the clear specication of a local problem if their work is to achieve realism. Clearly, adequate explanations for reproductive strategies in a particular taxonomic group are inextricably wed to specic ecological patterns and processes that have an unavoidable historical component. Useful generalization is what is left after the impact of historical effects has been removed. It is hard to see how we could arrive at this understanding by rst claiming that history has had no important impact. In this example, the basic focal level is specied by our assumptions about the basic units of selection and evolution. Most theories dealing with the distribution of sexuality are based on the genes-eye-view (Bell 1982), even though the Darwinian theory of evolution is based on differential survival and reproduction of individuals. However, we are convinced by Schmalgauzens (1938) persuasive argument that individuals are integrated wholes that cannot be torn into parts. It is also relevant that, in the genes-eye-view, the term phenotype is often used as interchangeable with genotype (modeled at the level of one or a few loci) as if all individuals of a same phenotype were genetically identical, and each genotype would produce a unique phenotype, when both propositions are demonstrably false. Use of such simplications

227 may commonly lead to a failure of gene-based theories to apply at higher levels of integration. In general, workers interested in explaining the evolution of sex have followed the approach of comparing amphimictic and apomictic reproduction by contrasting, through simulation or thought experiments, the success of sexual populations or species with those that are asexual. However, apomictic species (or races) are pragmatic taxonomic units which lump together clones that are morphologically similar and treat them as equivalent to sexually reproducing species (races). Setting these types as equivalent may lead to biased conclusions about the relative success of the two strategies. This follows because amphimictic populations are connected through reshufing and recombination of genetic material of interbreeding parents. In electrical jargon, they are connected in parallel. In contrast, clonally reproducing lineages are reproductively isolated, and thus more like multiple series. A system built on series connection is vulnerable in the sense that a single failure in the sequence will disrupt the whole system. A failure in a system with parallel connection is less fatal, because the message can proceed along alternative lines. Thus for sexually reproducing taxa, populations are inherently redundant, but this redundancy provides insurance against vagaries of the environment. If the above reasoning is accepted, generalizations based on comparisons of amphimictic and apomictic units should start by contrasting amphimictic populations with apomictic clones that are genetically more or less identical. Janzen (1977) has called such clones of apomictic organisms evolutionary individuals (EI). In contrast, some formal models (e.g., Maynard Smith 1978) have compared the success of a large number of specialized clones with that of one single amphimictic population so that the focal level for apomicts has been elevated to the focal population level for amphimicts. Of course, an EI is not identical with an individual, because the larger the number of replicate genomes of an EI, the smaller the role of chance in the exclusion of an EI. But once an apomictic lineage (genotype) is lost, it cannot be reconstituted through recombination as is the case in amphimictic populations. The importance of this difference in focal levels affects also scale and grain, central concepts in specifying ecological constraints. Because apomicts have fewer constraints against polyploidy than do sexually reproducing species, they can evolve increased heterozygosity through polyploidy and mutation (Bell 1982). The advantages of apomixis, then, might be sought in situations where high heterozygosity is advantageous. Heterozygosity, which is often associated with genetic homeostasis and heterosis (Lerner 1954) should be advantageous in extreme physical environments where organisms are affected by adversity selection (Greenslade 1983;

228 Southwood 1977, 1988). However, there are important reasons not to generalize that apomictic populations are universally superior to sexual ones under adversity selection. The advantages depend upon the grain of the environment. Depending on whether the environment is ne- or coarsegrained, the most t population is either one superior phenotype or a mixture of specialized phenotypes adapted to the relative frequencies of alternative environmental states (Levins 1968). It follows that the relative successes of apomicts and amphimicts may change with the amplitude and scale of the environmental changes relative to the life time of single individuals. Depending on the time scales of the alternating states and the life spans and generation lengths of the organisms, each apomictic lineage (multiple series) may go extinct in turn, but sexual populations (parallel series) respond more slowly to negative selection and are likely to survive longer in a relatively hard, coarse-grained environment. It follows that expansion of time scale may promote the level of signicant evolutionary process to that of species selection. Usually when reproductive modes are compared, the only initiating conditions considered are the cytological factors which determine the mode. However, different modes differ in evolutionary potential, and this may be associated with unexpected lower-level constraints. For example, apomicts are considered poor competitors, but good r-strategists of suitable earlysuccession habitats, partly because they may allegedly double the number of female offspring (all else being equal). However, population growth rate, an important aspect of r-strategies, is increased more easily by increasing the number of generations per season than by increasing the number of offspring per female, and so one should expect a high proportion of multivoltine species among apomicts. Paradoxically, apomictic insects tend to be univoltine, usually with a one- to three-year life cycle. Vepslinen and Jrvinen (1979) suggested that this paradox ows from the mechanism of diapause determination in insects. In general, facultatively multivoltine populations use an environmental cue such as day length to shut off reproduction and to stimulate metabolic preparations for a period without reproductive activity (usually winter in temperate latitudes). Because the value of the cue at the switchpoint (e.g., critical day length) changes with latitude, expansion from the source of origin of apomicts may be constrained by limited genetic variation for diapause determination. In other words, the ecological boundary conditions change with latitude, so that the mechanism of diapause determination works against dispersal of potentially multivoltine apomicts to new geographical areas. Apomictic insects are likely to belong to taxa that, for phylogenetic or ecological reasons, are obligatorily univoltine. Cyclically parthenogenetic organisms, on the other hand, are not evolution-

229 arily constrained by diapause determination. For example, both aphids and daphnids are known to be geographically wide-spread and highly multivoltine when the climate is favorable. The above examples illustrate that useful generalizations are likely to be built only with knowledge and understanding of biological details. Bold generalizations trying to simplify the whole universe to the level of a single, focused theory, by rst excluding the irrelevant details are likely to fail in many situations. It is not fatal if such failure inspires us to look more closely at the details. However, biologists do not receive much credit for failed theories and there is tremendous pressure to explain away or ignore failures. General theories are often the province of a majority of biologists who work on a small proportion of species. In summary, we advocate an explicit strategy of bottom-up pluralism. Instead of looking for single universal theories to explain the great variety of life, we suggest that it is more protable to develop general frameworks for explaining the evolution or ecological signicance of phenomena observable in nature and involving particular organisms. Such a framework should clearly specify focal levels (e.g., an apomictic clone belongs to a lower level of hierarchy than does an amphimictic population). Focal levels should be associated with the appropriate lower-level initiating conditions such as cytological constraints, and possible secondary constraints such as diapause determination, as in the example above. The framework should also outline the relevant kinds of upper-level ecological constraints that should be considered in providing explanations at each focal level (Figure 1). In other words, we argue that meaningful generality lies in the approach used to solve the problem, and that efforts to nd simple, universal theories, which are unbounded by copious contingency and yet satisfactorily predict characteristics of all groups of organisms, are doomed to failure.

The problem of scale and effective generalization Much useful research starts with a desire to explain a pattern and/or process observed in a specic natural system. We then seek to establish connections to either an upper-level pattern or a lower-level process, and in so doing test a set of hypotheses most of which derive at least in part from the general explanatory frameworks or paradigms of the day. Results of these initial tests usually lead us to explore the system in further detail by testing new sets of hypotheses, based on an interaction of our results and the explanatory framework. In doing this, we travel back and forth between levels to ensure that the connections are sound.

230

Figure 1. A schematic view of a research framework (stippled area) as embedded in the broader ecological and evolutionary sphere discussed in our example 4. Most ecological research is done by constructing and testing specic models of cause and effect relationships that describe how various processes produce patterns observed at the focal level of our interest. Within this overall framework, processes are driven by particular lower-level initiating conditions, which in our example include cytological mechanisms that determine mode of reproduction. Patterns are constrained by upper-level boundary conditions, represented by frequency of resource limitation that inuences the outcome of competitive interactions of apomictic and amphimictic lines. Frequently biologists wish to generalize the explanatory power of these frameworks over a number of taxa subject to similar lower-level initiating and upper-level boundary conditions. However, research frameworks are embedded in a matrix of interactions here partitioned among external factors, internal factors and historical factors. Each of these categories of inuence [accessory conditions of Taylor (1989)] may affect the focal pattern directly or affect it indirectly through the lower- and upper-level constraints here represented by black arrows. Robust generalizations depend critically on attention to these accessory conditions, the effects of which vary conspicuously among taxa, but which are generally controlled in specic experimental tests. Generalizations that neglect the inuence of these accessory conditions, which are not controlled in real-world situations, are bound to fail. We also note that the focal level of any particular research may provide initiating conditions for upper-level processes or also bound patterns at adjacent lower levels.

231 When shifting between levels ecologists must be especially aware of scale problems. This has been elaborated by Haila (1990) in studies of the power of the equilibrium theory of island biogeography (MacArthur and Wilson 1967). The general processes invoked in this theory should apply to a wide variety of islands, but successful application of the theory to particular situations depends crucially on choosing the correct scale. For example, for birds with migratory movements and high dispersal capacities, the geographical scale and isolation of the land archipelago of the Baltic Sea is too small for the results of equilibrium processes to be realized; any tendency toward equilibrium is constantly upset by the movement of birds. On the other hand, application of the theory to smaller organisms with much lower dispersal rates (e.g., carabid beetles) may be problematic because of relatively fast uplift in the Baltic basin (measured on an ecological time scale) which steadily increases the land area and alters the mix of habitats available. Taken as an explanatory framework, the theory has been an effective research tool in the eld of evolutionary ecology, pointing out the linkage of important factors affecting insular communities and encouraging their study. But if taken as a strict explanatory model of how the world works, the theory is generally found wanting the denition of what an islandis depends on the scales of the patterns and processes studied. Although most biological systems are continuous, it is convenient to consider the problem of scale in relation to biological hierarchies. Salthe (1985) has classied the living world along two hierarchies, the genealogical one emphasizing units of replication, and the ecological hierarchy emphasizing the organization of energy ow. The basic levels of these hierarchies are, respectively, genes genotypes/phenotypes demes species lineages, and molecules cells organisms populations ecosystems biosphere. Depending on the goals of the study, interstitial levels may be added, and other hierarchies may be outlined (e.g., Allen and Starr 1982; ONeill et al. 1986). But it is important to note that the boundaries between the hierarchies are set by the human mind (as may be some of the levels, e.g., the deme is more a theoretical concept than a real entity), and interactions between the hierarchies are manifold (Eldredge 1985; Liem 1990). To illustrate this, we revisit the problem of sex in metazoans. The study starts from observations: some living systems reproduce sexually, other ones asexually. This observation produces the question what, what kind of alternative reproductive modes do we have; this automatically calls for specication, achieved by collection of data about specic animals. The focal level of study is on the organismic and especially the population/clone level. With progress of study, connections in ecological, geographical and historical dimensions start to emerge, and simultaneously the focal level

232 of what tends to be elevated, and generalizations are made about what we observe. Because we are interested in alternative strategies, generalizations that are especially useful in directing the course of studies tend to be comparative. For example, whether it is true or not, the generalization The proportion of apomictic species increases with distance from the equator is useful, because it is testable by suitable data. If the pattern is real, we start to ask questions how it arose and why this particular evolutionary pathway was followed; the focus is on initiating conditions and on ecological boundary conditions, respectively. The data that are needed are, again, specic by nature. To understand alternative reproductive modes, one needs to know about the initiating conditions that are created at the cell level: details of cell division. It is evident that our new focus means that we traverse the boundary between Salthes two hierarchies, to study meiosis and mitosis and related processes, to learn and understand what kind of geno- and phenotypes can be produced in alternative reproductive modes. When we know how these are produced, we are able to understand more about that which we observed while generalizing our description at the level of populations and ecosystems. In the case of this example, the ultimate question for evolutionary biologists is why does a pattern exist. That question makes sense only after we have a sound set of descriptions of alternative reproductive strategies, and materially and theoretically justied generalizations about them. The basic focal level becomes again that of populations. Useful knowledge about initiating conditions may be derived from studies of phenotypic attributes and an understanding of phylogeny. Boundary conditions may be specied, by comparing specic reproductive modes in different ecological contexts, as integral parts of whole life-history strategies. But it should already be clear that most biological phenomena have many dimensions and will resist understanding as long as knowledge is restricted to one or a few levels of the biological hierarchies (Salthe 1985). Thus process or pattern that we wish to study denes our level of focus. The set of potential and actual states at the focal level interacts with conditions at the contiguous lower and upper levels of organization, through sets of many-to-one and one-to-many connections (Levins and Lewontin 1985). The number of initiating conditions and their permutations at the lower level dene the potential states at the focal level, whereas the actual state is constrained by the upper-level boundary conditions (see also Buss 1987). Because the focal level itself regulates, to some extent, actual conguration of states at the next-lower level, dynamic changes of the system at the focal level will affect the degrees of freedom for initiating conditions, and this dynamic is critical for the sequence of states observed. For example, let us return to

233 Figure 1 of our example 4. Given the potentials generated by the lowerlevel conditions and the constraints set by the upper-level factors, specic outcomes are realized at the focal level. One of the focal-level outcomes is a specic population-genetic structure that sets boundaries for the diversity of genotypes generated at the lower level on the basis of genetic and cytological mechanisms, specic to amphi- vs apomictic systems. Also, the focal-level patterns may act as initiating conditions for processes at the upper level. For example, the populations under study consume resources, and also affect accessory conditions in many possible ways (e.g., by affecting the population dynamics of predators controlling the focal-level processes). Salthe (1985) refers to these adjacent levels as the basic triadic system. Because transmission of information attenuates steeply with the distance between levels, Salthe suggests that an understanding of the triadic structure is generally sufcient for a minimal description of any complex diachronic system. The scale, or extent of spatio-temporal variation, of things at lower levels of nested hierarchies is smaller than the scale at upper levels. Complexity increases with the rank of the level in nested hierarchies because higher levels are a composite of all lower levels plus their interactions. To escape the dilemma of dealing with daunting complexity at higher levels, ecologists usually bound their generalizations and thus, restrict explanations to certain sets of lower-level parameters thus decreasing the domain of the explanation at the higher level. We believe that generalizations become increasingly sensitive to boundary conditions as we move upscale because the number of potential causal routes generating any pattern increases explosively reecting the many to one connections from lower levels. The same is true for scale expansions at the focal level, both in space and time, as these will also expand the complexity of constraints on the contiguous upper level. Thus the amount of generality achievable varies in the opposite direction to levels of organization, scale and complexity. However, strictly empirical generalizations that average out variability at lower levels may hold. The problem of generalization is to relate effectively the initiating and boundary conditions to the focal level in consistent terms as in examples 1 and 4 above. Intuitively we dene the right scale as the optimally manageable scale at the focal level, allowing formulation of explanations and testable hypotheses, and experimental control over contiguous lower- and/or higherlevel conditions. As Salthe (1985, p. 96) put it: Experimental design is often the art of nding only what one is looking for (that is why it is often associated with the strategy of conrmation rather than that of falsication). When information about boundary conditions is increased, predictability grows on the scale of interest. However, predictability does not in itself confer generality, because generality is dependent on our understanding of the relevant

234 complexity, not knowledge of specic outcomes. Taylor (1989) raises the important point that robust generality is evasive and not a necessary outcome for every scientic paper; there must be room to explain our conceptual frameworks and their connections without covering up their lack of correspondence with empirical reality. When the focal level is reduced, boundary conditions tend to become more restrictive allowing fewer possibilities, and the predictions can be more general because of this reduced complexity. The scale and complexity of the biological phenomenon to be explained determines the appropriate scales for observations and experiments to illuminate reciprocally initiating and regulating conditions. Through iterations of this process, in work with different systems, we test, modify and bound our higher-level generalizations, while being somewhat unconcerned about whether these generalizations are strictly true. When a robust and general explanatory pattern emerges from this process, we are converging on the truth, truth being appropriately dened as the intersection of independent lies (Levins 1966).

Acknowledgements We thank Yrj Haila, Christian Klingenberg, Jari Niemel and Peter Taylor for helpful discussion and comments on the manuscript, and J. S. Scott for drafting the gure. JRS thanks T. R. E. Southwood for several detailed discussions on the habitat templet. This manuscript was conceived and largely completed in 1991 during a years visit by KV to the Department of Entomology, University of Alberta which was supported by an NSERC International Scientic Exchange Grant and a grant from the Academy of Finland for which we are grateful. The work was also supported by an NSERC Operating Grant to JRS.

Note
1 All this said, and given the known large variety of mating behavior of waterstriders (Spence

and Andersen 1994), it is disappointing to see that a generalized (type one) mating sequence of striders (Rowe et al. 1994; Arnqvist 1997) includes premating struggle, guarding, and postmating struggle leading to dislodgment of the male, but not oviposition. This generalization is strongly dissonant with the theoretical value of guarding, which should maximize the probability that the guarding male will father the offspring (Yamamura 1987). Such discrepancy between theory and experimental results is alarming, especially if no alternative explanation for guarding is given and no reference is made to measures taken to conrm the lack of oviposition. When mating experiments are done in spacious enclosures in the natural habitat of waterstriders, females of two common Gerris species are found to oviposit while guarded by

235
the male (Vepslinen and Spence, unpublished). Such forced generalizations are maintained even in spite of published evidence to the contrary. Vepslinen and Savolainen (1995) showed how termination of mating in a waterstrider species depends on the individual experiences of males and females, so that depending on the experimental treatment, the vast majority of the matings can be terminated by the male, without preceding struggle or guarding. After arguing repeatedly against acceptance of the Vepslinen and Savolainen manuscript as an anonymous reviewer for three highly respected journals in a row, Arnqvist (1997) eventually included our (nally published) nding in his (otherwise unchanged) generalized mating sequence as the footnote: male reported to terminate mating occasionally. It is difcult to see how such oversimplication could help to understand highly exible behaviors. Generalizing one tactic at the cost of all alternative tactics will, by denition, delay synthesis leading to an understanding of mating strategies. Overprotection of faulty generalizations is encouraged by a peer-review system in which perpetrators of generalizations under question gure largely in decisions about whether opposing evidence will be communicated, especially when irresponsible reviewers are able to dodge accountability through anonymity.

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