The Epic of Evolution is a course taught at Northern Arizona University. It engages the task of formulating a new epic myth that is based on the physical, natural, social, and cultural sciences. It aims to serve the need of providing meaning for human living in the vast and complex universe that the sciences now depict for us. It is an interdisciplinary effort in an academic setting that is often divided by specializations; it focuses on values in a climate (...) of relativism; and it concentrates on an enterprise for which there are few textbooks at hand. The course is presented in three segments: the cosmos before humans appeared, the human phenomenon, and scenarios for the future of evolution. (shrink)

Complexity is pleased to announce the installment of Prof Hiroki Sayama as its new Chief Editor. In this Editorial, Prof Sayama describes his feelings about his recent appointment, discusses some of the journal’s journey and relevance to current issues, and shares his vision and aspirations for its future.

The study of culture from an evolutionary perspective has been slowed by resistance from some quarters of anthropology, a poor appreciation of the fidelity of cultural transmission, and misunderstandings about human intentionality. (Published Online November 9 2006).

In the scientific literature on religious evolution, two competing theories appeal to group selection to explain the relationship between religious belief and altruism, or costly, prosocial behavior. Both theories agree that group selection plays an important role in cultural evolution, affecting psychological traits that individuals acquire through social learning. They disagree, however, about whether group selection has also played a role in genetic evolution, affecting traits that are inherited genetically. Recently, Jonathan Haidt has defended the most (...) fully developed account based on genetic group selection, and I argue here that problems with this account reveal good reasons to doubt that genetic group selection has played any important role in human evolution at all. Thus, considering the role of group selection in religious evolution is important not just because of what it reveals about religious psychology and religious evolution, but also because of what it reveals about the role of group selection in human evolution more generally. (shrink)

We hypothesize that heritable epigenetic changes can affect rates of fitness increase as well as patterns of genotypic and phenotypic change during adaptation. In particular, we suggest that when natural selection acts on pure epigenetic variation in addition to genetic variation, populations adapt faster, and adaptive phenotypes can arise before any genetic changes. This may make it difficult to reconcile the timing of adaptive events detected using conventional population genetics tools based on DNA sequence data with environmental drivers (...) of adaptation, such as changes in climate. Epigenetic modifications are frequently associated with somatic cell differentiation, but recently epigenetic changes have been found that can be transmitted over many generations. Here, we show how the interplay of these heritable epigenetic changes with genetic changes can affect adaptive evolution, and how epigenetic changes affect the signature of selection in the genetic record. (shrink)

Burt's target article oddly misses the important intellectual contribution of sociogenomics to our understanding of genetic evolution in contemporary human populations. Although social scientists' immediate research agendas are often not evolutionary in nature, I call for a better appreciation of the role of sociogenomics in answering important evolutionary questions.

The article proposes to further develop the ideas of the Extended Evolutionary Synthesis by including into evolutionary research an analysis of phenomena that occur above the organismal level. We demonstrate that the current Extended Synthesis is focused more on individual traits (genetically or non-genetically inherited) and less on community system traits (synergetic/organizational traits) that characterize transgenerational biological, ecological, social, and cultural systems. In this regard, we will consider various communities that are made up of interacting populations, and for which the (...) individual members can belong to the same or to different species. Examples of communities include biofilms, ant colonies, symbiotic associations resulting in holobiont formation, and human societies. The proposed model of evolution at the level of communities revises classic theorizing on the major transitions in evolution by analyzing the interplay between community/social traits and individual traits, and how this brings forth ideas of top-down regulations of bottom-up evolutionary processes (collaboration of downward and upward causation). The work demonstrates that such interplay also includes reticulate interactions and reticulate causation. In this regard, we exemplify how community systems provide various non-genetic ‘scaffoldings’, ‘constraints’, and ‘affordances’ for individual and sociocultural evolutionary development. Such research complements prevailing models that focus on the vertical transmission of heritable information, from parent to offspring, with research that instead focusses on horizontal, oblique and even reverse information transmission, going from offspring to parent. We call this reversed information transfer the ‘offspring effect’ to contrast it from the ‘parental effect’. We argue that the proposed approach to inheritance is effective for modelling cumulative and distributed developmental process and for explaining the biological origins and evolution of language. (shrink)

Darwinian evolutionary theory has two key terms, variations and biological selection, which finally lead to survival of the fittest variant. With the rise of molecular genetics, variations were explained as results of error replications out of the genetic master templates. For more than half a century, it has been accepted that new genetic information is mostly derived from random error-based events. But the error replication narrative has problems explaining the sudden emergence of new species, new phenotypic traits, and (...) genome innovations as a sudden single event. Meanwhile, it is recognized that errors cannot explain the evolution of genetic information, genetic novelty, and complexity. Now, empirical evidence establishes the crucial role of non-random genetic content editors, such as viruses, diversity generating retroelements, and other RNA networks, to produce new genetic information, complex regulatory control, inheritance vectors, genetic identity, immunity, new sequence space, evolution of complex organisms, and evolutionary transitions. (shrink)

This paper explores how an evolutionary process can produce systems that learn. A general framework for the evolution of learning is outlined, and is applied to the task of evolving mechanisms suitable for supervised learning in single-layer neural networks. Dynamic properties of a network’s information-processing capacity are encoded genetically, and these properties are subjected to selective pressure based on their success in producing adaptive behavior in diverse environments. As a result of selection and genetic recombination, various successful learning (...) mechanisms evolve, including the well-known delta rule. The effect of environmental diversity on the evolution of learning is investigated, and the role of different kinds of emergent phenomena in genetic and connectionist systems is discussed. (shrink)

Ideas about heredity and evolution are undergoing a revolutionary change. New findings in molecular biology challenge the gene-centered version of Darwinian theory according to which adaptation occurs only through natural selection of chance DNA variations. In Evolution in Four Dimensions, Eva Jablonka and Marion Lamb argue that there is more to heredity than genes. They trace four "dimensions" in evolution -- four inheritance systems that play a role in evolution: genetic, epigenetic, behavioral, and symbolic. These (...) systems, they argue, can all provide variations on which natural selection can act. Evolution in Four Dimensions offers a richer, more complex view of evolution than the gene-based, one-dimensional view held by many today. The new synthesis advanced by Jablonka and Lamb makes clear that induced and acquired changes also play a role in evolution.After discussing each of the four inheritance systems in detail, Jablonka and Lamb "put Humpty Dumpty together again" by showing how all of these systems interact. They consider how each may have originated and guided evolutionary history and they discuss the social and philosophical implications of the four-dimensional view of evolution. Each chapter ends with a dialogue in which the authors engage the contrarieties of the fictional "I.M.," or Ifcha Mistabra -- Aramaic for "the opposite conjecture" -- refining their arguments against I.M.'s vigorous counterarguments. The lucid and accessible text is accompanied by artist-physician Anna Zeligowski's lively drawings, which humorously and effectively illustrate the authors' points. (shrink)

Behavioral genetics and cultural evolution have both revolutionized our understanding of human behavior – largely independent of each other. Here, we reconcile these two fields under a dual inheritance framework, offering a more nuanced understanding of the interaction between genes and culture. Going beyond typical analyses of gene–environment interactions, we describe the cultural dynamics that shape these interactions by shaping the environment and population structure. A cultural evolutionary approach can explain, for example, how factors such as rates of innovation (...) and diffusion, density of cultural subgroups, and tolerance for behavioral diversity impact heritability estimates, thus yielding predictions for different social contexts. Moreover, when cumulative culture functionally overlaps with genes, genetic effects become masked, unmasked, or even reversed, and the causal effects of an identified gene become confounded with features of the cultural environment. The manner of confounding is specific to a particular society at a particular time, but a WEIRD (western, educated, industrialized, rich, and democratic) sampling problem obscures this boundedness. Cultural evolutionary dynamics are typically missing from models of gene-to-phenotype causality, hindering generalizability of genetic effects across societies and across time. We lay out a reconciled framework and use it to predict the ways in which heritability should differ between societies, between socioeconomic levels, and other groupings within some societies but not others, and over the life course. An integrated cultural evolutionary behavioral genetic approach cuts through the nature–nurture debate and helps resolve controversies in topics such as IQ. (shrink)

Our understanding of language, its origins and subsequent evolution, is shaped not only by data and theories from the language sciences, but also fundamentally by the biological sciences. Recent developments in genetics and evolutionary theory offer both very strong constraints on what scenarios of language evolution are possible and probable, but also offer exciting opportunities for understanding otherwise puzzling phenomena. Due to the intrinsic breathtaking rate of advancement in these fields, and the complexity, subtlety, and sometimes apparent non-intuitiveness (...) of the phenomena discovered, some of these recent developments have either being completely missed by language scientists or misperceived and misrepresented. In this short paper, we offer an update on some of these findings and theoretical developments through a selection of illustrative examples and discussions that cast new light on current debates in the language sciences. The main message of our paper is that life is much more complex and nuanced than anybody could have predicted even a few decades ago, and that we need to be flexible in our theorizing instead of embracing a priori dogmas and trying to patch paradigms that are no longer satisfactory. (shrink)

The biogenesis of RNAs and proteins is a threat to the cell. Indeed, the act of transcription and nascent RNAs challenge DNA stability. Both RNAs and nascent proteins can also initiate the formation of toxic aggregates because of their physicochemical properties. In reviewing the literature, I show that co-transcriptional and co-translational biophysical constraints can trigger DNA instability that in turn increases the likelihood that sequences that alleviate the constraints emerge over evolutionary time. These directed genetic variations rely on the (...) biogenesis of small RNAs that are transcribed directly from challenged DNA regions or processed from the transcripts that directly or indirectly generate constraints or aggregates. These small RNAs can then target the genomic regions from which they initially originate and increase the local mutation rate of the targeted loci. This mechanism is based on molecular pathways involved in anti-parasite genome defence systems, and implies that gene expression-related biophysical constraints represent a driving force of genome evolution. Are randomly generated mutations the only source of genetic variation during evolution? This paper describes the molecular mechanisms by which co-transcriptional and co-translational biophysical constraints trigger genetic variations in targeted-genomic sequences in an RNA-depending manner. This directed-mutational process that drives genome evolution is related to antiparasite genome defence systems. (shrink)

The paper describes the context and the origin of a particular debate that concerns the evolution of phenotypic plasticity. In 1965, British biologist A. D. Bradshaw proposed a widely cited model intended to explain the evolution of norms of reaction, based on his studies of plant populations. Bradshaw’s model went beyond the notion of the “adaptive norm of reaction” discussed before him by Dobzhansky and Schmalhausen by suggesting that “plasticity” the ability of a phenotype to be modified by (...) the environment should be genetically determined. To prove Bradshaw’s hypothesis, it became necessary for some authors to identify the pressures exerted by natural selection on phenotypic plasticity in particular traits, and thus to model its evolution. In this paper, I contrast two different views, based on quantitative genetic models, proposed in the mid-1980s: Russell Lande and Sara Via’s conception of phenotypic plasticity, which assumes that the evolution of plasticity is linked to the evolution of the plastic trait itself, and Samuel Scheiner and Richard Lyman’s view, which assumes that the evolution of plasticity is independent from the evolution of the trait. I show how the origin of this specific debate, and different assumptions about the evolution of phenotypic plasticity, depended on Bradshaw’s definition of plasticity and the context of quantitative genetics. (shrink)

The study of genetic developmental disorders originally seemed to hold the promise for those of a nativist persuasion of demonstrating pure dissociations between different cognitive functions, as well as the existence of innately specified modules in the brain and the direct mapping of mutated genes to specific cognitive-level outcomes. However, more recent research within a neuroconstructivist perspective has challenged this promise, arguing that earlier researchers lost sight of one fundamental explanatory factor in both the typical and atypical case: the (...) actual process of ontogenetic development. The paper is divided into three parts on evolution, genetics, and ontogeny. Each section starts by examining nativist claims about innateness and modularity of function, and subsequently evaluates them within a neuroconstructivist approach to human development. (shrink)

There are currently three major theories on the origin and evolution of the genetic code: the stereochemical theory, the coevolution theory, and the error-minimization theory. The first two assume that the genetic code originated respectively from chemical affinities and from metabolic relationships between codons and amino acids. The error-minimization theory maintains that in primitive systems the apparatus of protein synthesis was extremely prone to errors, and postulates that the genetic code evolved in order to minimize the (...) deleterious effects of the translation errors. This article describes a fourth theory which starts from the hypothesis that the ancestral genetic code was ambiguous and proposes that its evolution took place with a mechanism that systematically reduced its ambiguity and eventually removed it altogether. This proposal is distinct from the stereochemical and the coevolution theories because they do not contemplate any ambiguity in the genetic code, and it is distinct from the error-minimization theory because ambiguity-reduction is fundamentally different from error-minimization. The concept of ambiguity-reduction has been repeatedly mentioned in the scientific literature, but so far it has remained only an abstract possibility because no model has been proposed for its mechanism. Such a model is described in the present article and may be the first step in a new approach to the study of the evolution of the genetic code. (shrink)

Why do humans menstruate while most mammals do not? Here, we present our answer to this long‐debated question, arguing that (i) menstruation occurs as a mechanistic consequence of hormone‐induced differentiation of the endometrium (referred to as spontaneous decidualization, or SD); (ii) SD evolved because of maternal–fetal conflict; and (iii) SD evolved by genetic assimilation of the decidualization reaction, which is induced by the fetus in non‐menstruating species. The idea that menstruation occurs as a consequence of SD has been proposed (...) in the past, but here we present a novel hypothesis on how SD evolved. We argue that decidualization became genetically stabilized in menstruating lineages, allowing females to prepare for pregnancy without any signal from the fetus. We present three models for the evolution of SD by genetic assimilation, based on recent advances in our understanding of the mechanisms of endometrial differentiation and implantation. Testing these models will ultimately shed light on the evolutionary significance of menstruation, as well as on the etiology of human reproductive disorders like endometriosis and recurrent pregnancy loss. (shrink)

Here reprinted from the 1902 Macmillan edition.

The essays in this collection examine developments in three fundamental biological disciplines--embryology, evolutionary biology, and genetics--in conflict with each other for much of the twentieth century. They consider key methodological problems and the difficulty of overcoming them. Richard Burian interweaves historical appreciation of the settings within which scientists work, substantial knowledge of the biological problems at stake and the methodological and philosophical issues faced in integrating biological knowledge drawn from disparate sources.

The idea that development is the expression of information accumulated during evolution and that heredity is the transmission of this information is surprisingly hard to cash out in strict, scientific terms. This paper seeks to do so using the sense of information introduced by Francis Crick in his sequence hypothesis and central dogma of molecular biology. It focuses on Crick's idea of precise determination. This is analysed using an information-theoretic measure of causal specificity. This allows us to reconstruct some (...) of Crick's claims about information in transcription and translation. Crick's approach to information has natural extensions to non-coding regions of DNA, to epigenetic marks, and to the genetic or environmental upstream causes of those epigenetic marks. Epigenetic information cannot be reduced to genetic information. The existence of biological information in epigenetic and exogenetic factors is relevant to evolution as well as to development. (shrink)

There are many ways that biological theory can inform ethical discussions of genetic engineering and biomedical enhancement. In this essay, we highlight some of these potential contributions, and along the way provide a synthetic overview of the papers that comprise this special issue. We begin by comparing and contrasting genetic engineering with programs of selective breeding that led to the domestication of plants and animals, and we consider how genetic engineering differs from other contemporary biotechnologies such as (...) embryo selection. We go on to consider the implications of genetic engineering for human nature, human evolution, and persistence of the human species. Finally, we question whether genetic interventions warrant the extraordinary ethical scrutiny they are often given, and we show how the misleading “genetic blueprint” metaphor has imposed a faulty structure on the enhancement debate. We conclude by considering the nature of biological development and the sobering limits it places on what genetic engineering can reasonably hope to achieve. (shrink)

Many philosophers invoke the "wisdom of nature" in arguing for varying degrees of caution in the development and use of genetic enhancement technologies. Because they view natural selection as akin to a master engineer that creates functionally and morally optimal design, these authors tend to regard genetic intervention with suspicion. In Part II, we examine and ultimately reject the evolutionary assumptions that underlie the master engineer analogy (MEA). By highlighting the constraints on ordinary unassisted evolution, we show (...) how intentional genetic modification can overcome many of the natural impediments to the human good. Our contention is that genetic engineering offers a solution that is more eff icient, reliable, versatile, and morally palatable than the lumbering juggernaut of Darwinian evolution. In Part III, we evaluate a recent attempt to ground precautionary enhancement heuristics in adaptive etiology. Our problem with this approach is two-fold: first, it is based on the same "strong adaptationist" interpretation of evolution that motivates the flawed MEA, and second, the etiological concept of function on which it relies provides indirect and potentially misleading information about the likely consequences of genetic intervention. We offer instead enhancement criteria based on causal relationships in ontogeny. We conclude that rather than grounding a presumption against deliberate genetic modification, the causal structure of the living world gives us good moral reason to pursue it. (shrink)

There is growing interest in the evolutionary dynamics of molecular genetic pathways and networks, and the extent to which the molecular evolution of a gene depends on its position within a pathway or network, as well as over‐all network topology. Investigations on the relationships between network organization, topological architecture and evolutionary dynamics provide intriguing hints as to how networks evolve. Recent studies also suggest that genetic pathway and network structures may influence the action of evolutionary forces, and (...) may play a role in maintaining phenotypic robustness in organisms. BioEssays 26:479–484, 2004. © 2004 Wiley Periodicals, Inc. (shrink)

Collected for the first time in a single volume are essays which examine the developments in three fundamental biological disciplines - embryology, evolutionary biology, and genetics. These disciplines were in conflict for much of the twentieth century and the essays in this collection examine key methodological problems within these disciplines and the difficulties faced in overcoming the conflicts between them. Burian skilfully weaves together historical appreciation of the settings within which scientists work, substantial knowledge of the biological problems at stake (...) and the methodological and philosophical issues faced in integrating biological knowledge drawn from disparate sources. The final chapter describes what recent findings in development biology and genetics can tell us about the history and development of animals. Written in a clear, accessible style this collection should appeal to students and professionals in philosophy of science, and the philosophy and history of biology. (shrink)

This book presents a survey of the molecular basis for the genetic control of living organisms and their evolution. The authors consider four dimensions of control over what shapes life forms: genetic, epigenetic, behavioral, and symbolic/cultural. They pay particular attention to the epigenetic realm, and they defend a view recognizing the genetic incorporation of acquired characteristics – a neo-Lamarckian tack.

In their recent book, Elliott Sober and David Wilson (1998) argue that evolutionary biologists have wrongly regarded kinship as the exclusive means by which altruistic behavior can evolve, at the expense of other mechanisms. I argue that Sober and Wilson overlook certain genetical considerations which suggest that kinship is likely to be a more powerful means for generating complex altruistic adaptations than the alternative mechanisms they propose.

We extend previous work by modeling evolution of communication using a spatialized genetic algorithm which recombines strategies purely locally. Here cellular automata are used as a spatialized environment in which individuals gain points by capturing drifting food items and are 'harmed' if they fail to hide from migrating predators. Our individuals are capable of making one of two arbitrary sounds, heard only locally by their immediate neighbors. They can respond to sounds from their neighbors by opening their mouths (...) or by hiding. By opening their mouths in the presence of food they maximize gains; by hiding when a predator is present they minimize losses. We consider the result a 'natural' template for benefits from communication; unlike a range of other studies, it is here only the recipient of communicated information that immediately benefits. (shrink)

While prior models of the evolution of altruism have assumed that organisms reproduce asexually, this paper presents a model of the evolution of altruism for sexually reproducing organisms using Hardy–Weinberg dynamics. In this model, the presence of reciprocal altruists allows the population to evolve to a stable polymorphic population where the majority of organisms are altruistic. Further, adding stochasticity leads to even larger numbers of altruists, while adding stochasticity to an analogous asexual model leads to more selfish organisms. (...) The contrast between these outcomes demonstrates why it may be important to pay attention to the underlying genetics of a population. (shrink)

The role of genetic relatedness in social evolution has recently come under critical attention. These arguments are here critically analyzed, both theoretically and empirically. It is argued that when the conceptual structure of the theory of natural selection is carefully taken into account, genetic relatedness can be seen to play an indispensable role in the evolution of both facultative and advanced eusociality. Although reviewing the empirical evidence concerning the evolution of eusociality reveals that relatedness does (...) not play a role in the initial appearance of helper phenotypes, this follows simply from the fact that natural selection – of which relatedness is a necessary component – does not play a causal role in the origin of any traits. Further, separating two logically distinct elements of causal explanation – necessity and sufficiency – explains why the debate lingers on: although relatedness plays a necessary role in the evolution of helping and advanced eusociality, relatedness alone is not sufficient for their appearance. Therefore, if the relatedness variable in a given data set is held at a uniformly high value, then it indeed may turn out that other factors occupy a more prominent role. However, this does not change the fact that high relatedness functions as a necessary background condition for the evolution of advanced eusociality. (shrink)

An analogy is drawn between the processes of human language evolution and the ongoing discoveries concerning how the human genome is constructed. Mutational evolution may be thought of in linguistic terms as an alternation in the genetic code following morphemic substitutions, deletions or additions. This may be termed an endosemiotic analysis where semiotic processes may be found at the biochemical level of the genome. Hence, owing to these genetic changes, phenotypic alterations in the morphology of the (...) organism create those evolutionary changes seen in the development of the phyla in the paleontological record. We are now witnessing the inclusion of the genome of all species both plant and animal within our material culture as we begin to modify these genotypes with recombinant DNA/rna technology. The coevolution of language and the genetic code may then occur as we begin to more thoroughly understand these interrelated evolutionary processes. (shrink)

The Segregation Distorter (SD) system of Drosophila melanogaster is one the best‐characterized meiotic drive complexes known. SD gains an unfair transmission advantage through heterozygous SD/SD+ males by incapacitating SD+‐bearing spermatids so that virtually all progeny inherit SD. Segregation distorter (Sd), the primary distorting locus in the SD complex, is a truncated duplication of the RanGAP gene, a major regulator of the small GTPase Ran, which has several functions including the maintenance of the nucleocytoplasmic RanGTP concentration gradient that mediates nuclear transport. (...) The truncated Sd‐RanGAP protein is enzymatically active but mislocalizes to the nucleus where it somehow causes distortion. Here I present data consistent with the idea that wild‐type RanGAP, and possibly other loci able to influence the RanGTP gradient, has been caught up in an ancient genetic conflict that predates the SD complex. The legacy of this conflict could include the unexpectedly rapid evolution of nuclear transport‐related proteins, the accumulation of chromosomal inversions, the recruitment of gene duplications, and the turnover of repetitive sequences in the centric heterochromatin. BioEssays 29:386–391, 2007. © 2007 Wiley Periodicals, Inc. (shrink)

In addition to considerable debate in the recent evolutionary literature about the limits of the Modern Synthesis of the 1930s and 1940s, there has also been theoretical and empirical interest in a variety of new and not so new concepts such as phenotypic plasticity, genetic assimilation and phenotypic accommodation. Here we consider examples of the arguments and counter- arguments that have shaped this discussion. We suggest that much of the controversy hinges on several misunderstandings, including unwarranted fears of a (...) general attempt at overthrowing the Modern Synthesis paradigm, and some fundamental conceptual confusion about the proper roles of phenotypic plasticity and natural selection within evolutionary theory. (shrink)

Although features of the dentition figure prominently in discussions of early hominid phylogeny, remarkably little is known of the developmental basis of the variations in occlusal morphology and dental proportions that are observed among taxa. Recent experiments on tooth development in mice have identified some of the genes involved in dental patterning and the control of tooth specification. These findings provide valuable new insight into dental evolution and underscore the strong developmental links that exist among the teeth and the (...) jaws and cranium. The latter has important implications for cladistic studies that traditionally consider features of the skull independently from the dentition. BioEssays 23:481–493, 2001. © 2001 John Wiley & Sons, Inc. (shrink)

We advocate for an integrative long-term perspective on time, noting that culture changes on timescales amenable to behavioral genetic study with appropriate design and modeling extensions. We note the need for replications of behavioral genetic studies to examine model invariance across long-term timescales, which would afford examination of specified as well as unspecified cultural moderators of behavioral genetic effects.

Language and life history can be related functionally through the study of human ontogeny, thus usefully informing our understanding of several unique aspects of the evolution of species. The operational principles outlined by Locke & Bogin (L&B) demonstrate that the present can provide a useful framework for understanding the past.

I define a concept of causal probability and apply it to questions about the role of probability in evolutionary processes. Causal probability is defined in terms of manipulation of patterns in empirical outcomes by manipulating properties that realize objective probabilities. The concept of causal probability allows us see how probabilities characterized by different interpretations of probability can share a similar causal character, and does so in such way as to allow new inferences about relationships between probabilities realized in different chance (...) setups. I clarify relations between probabilities and properties defined in terms of them, and argue that certain widespread uses of computer simulations in evolutionary biology show that many probabilities relevant to evolutionary outcomes are causal probabilities. This supports the claim that higher-level properties such as biological fitness and processes such as natural selection are causal properties and processes, contrary to what some authors have argued. (shrink)