This paper aims to illustrate one of the primary goals of the philosophy of biology⎯namely, the examination of central concepts in biological theory and practice⎯through an analysis of the concepts of population and metapopulation in evolutionary biology and ecology. I will first provide a brief background for my analysis, followed by a characterization of my proposed concepts: the causal interactionist concepts of population and metapopulation. I will then illustrate how the concepts apply to six cases that differ in their population (...) structure; this analysis will also serve to flesh out and defend the concepts a bit more. Finally, I will respond to some possible questions that my analysis may have raised and then conclude briefly. (shrink)

In a previous study, using experimental metapopulations of the flour beetle, Tribolium castaneum, we investigated phase III of Wright's shifting balance process (Wade and Griesemer 1998). We experimentally modeled migration of varying amounts from demes of high mean fitness into demes of lower mean fitness (as in Wright's characterization of phase III) as well as the reciprocal (the opposite of phase III). We estimated the meta-populational heritability for this level of selection by regression of offspring deme means on the (...) weighted parental deme means.Here we develop a Punnett Square representation of the inheritance of the group mean to place our empirical findings in a conceptual context similar to Mendelian inheritance of individual traits. The comparison of Punnett Squares for individual and group inheritance shows how the latter concept can be rigorously defined and extended despite the lack of explicitly formulated, simple Mendelian laws of inheritance at the group level. Whereas Wright's phase III combines both interdemic selection and meta-populational inheritance, our formulation separates the issue of meta-populational heritability from that of interdemic selection. We use this conceptual context to discuss the controversies over the levels of selection and the units of inheritance. (shrink)

The financial risk information diffuses through various kinds of social networks, such as Twitter and Facebook. Individuals transmit the financial risk information which can migrate among different platforms or forums. In this paper, we propose a financial risk information spreading model on metapopulation networks. The subpopulation represents a platform or forum, and individuals migrate among them to transmit the information. We use a discrete-time Markov chain approach to describe the spreading dynamics’ evolution and deduce the outbreak threshold point. We perform (...) numerical simulation on artificial networks and discover that the financial risk information can be promoted once increasing the information transmission probability and active subpopulation fraction. The weight variance and migration probability cannot significantly affect the financial risk spreading size. The discrete-time Markov chain approach can reasonably predict the above phenomena. (shrink)

In coarse-grained environments specialists are generally predicted to dominate. Empirically, however, coexistence with generalists is often observed. We present a simple, but previously unrecognized, mechanism for coexistence of a habitat generalist and a number of habitat specialist species. In our model all species have a metapopulation structure in a landscape consisting of patches of different habitat types, governed by local extinction and colonization. Each specialist is limited to its specific type of habitat. The generalist can use more types of habitat, (...) has a lower local competitive ability but can exploit patches left open by the specialists. Our modeling shows that coexistence is easily possible. The mechanism amounts to a colonization/competition trade-off at the landscape level, where the colonization advantage of the inferior competitor does not arise from a higher colonization rate but from its ability to use more types of habitat. Habitat availability has to be intermediate: when there are few patches of each habitat, only the generalist is able to maintain itself and when there are many patches, high propagule pressure of the specialists excludes the generalist. Habitat selection or temporal variations in relative habitat quality are not necessary for coexistence. Increased niche-width, colonization rate or local competitive ability of the generalist enhances its performance compared to the specialists. Various types of habitat degradation favour generalism. When able to use a broad range of habitats, generalists can generate so much propagule pressure that only a low level of local competitive ability is needed to globally exclude the specialists. Hence, in a reversal of the original problem, the question is why there are so many specialist metapopulations? (shrink)

We present a metapopulation dynamic model for the diffusively-coupled rock-paper-scissors game with mutation in scale-free hierarchical networks. We investigate how the RPS game changes by mutation in scale-free networks. Only the mutation from rock to scissors occurs with rate μ. In the network, a node represents a patch where the RPS game is performed. RPS individuals migrate among nodes by diffusion. The dynamics are represented by the reaction-diffusion equations with the recursion formula. We study where and how species coexist or (...) go extinct in the scale-free network. We numerically obtained the solutions for the metapopulation dynamics and derived the transition points. The results show that, with increasing mutation rate μ, the extinction of P species occurs and then the extinction of R species occurs, and finally only S species survives. Thus, the first and second dynamical phase transitions occur in the scale-free hierarchical network. We also show that the scaling law holds for the population dynamics which suggests that the transition points approach zero in the limit of infinite size. (shrink)

This paper examines the species problem in microbiology and its implications for the species problem more generally. Given the different meanings of ‘species’ in microbiology, the use of ‘species’ in biology is more multifarious and problematic than commonly recognized. So much so, that recent work in microbial systematics casts doubt on the existence of a prokaryote species category in nature. It also casts doubt on the existence of a general species category for all of life (one that includes both prokaryotes (...) and eukaryotes). Prokaryote biology also undermines recent attempts to save the species category, such as the suggestion that species are metapopulation lineages and the idea that ‘species’ is a family resemblance concept. (shrink)

The contextual approach is a prominent framework for thinking about group selection. Here, I highlight ambiguity about what the contextual approach is. Then, I discuss problematic entailments the contextual approach has for what processes count as group selection—entailments more troublesome than typically noted. However, Sober and Wilson’s version of the Price approach, which is the main alternative to the contextual approach, is problematic too: it leads to an underappreciated paradox called the vanishing selection problem and thereby generates the wrong qualitative (...) account of whether group selection is occurring in a certain family of cases. In response, I develop an account of group selection that can deal with the counterexamples to both the contextual approach and the Price approach. I then discuss the role that contextual analysis can continue to play in the discussion of individual fitness and metapopulation evolution. (shrink)

This paper proposes a new and testable view about the nature of species and other evolving lineages, according to which they are feedback systems. On this view, it is a mistake to think gene flow, niche sharing, and trait frequency similarities between populations are among variables that interact to cause some further downstream variable that distinguishes evolving lineages from each other, some sort of “species cohesion” for example. Instead, gene flow, niche sharing, similarities between populations, and other causal variables feed (...) into each other—instances of these at earlier times help cause instances of these same variables at later times. And any lineage-identifying cohesion just is the recurrence or cycling of these feedback relations within metapopulations over generations. Such cohesion can then be represented as variable M within multi-dimensional variable spaces, where values of M vary dynamically with the frequency and magnitude of feedback relations. Related conditions for being a species or other evolving lineage are then clarified. To argue for the development and testing of this view, the paper shows how it improves upon others. (shrink)

After having reconstructed a minimal biological characterisation of species, we endorse an “empirical approach” based on the idea that it is the peculiar evolutionary history of the species at issue—its peculiar origination process, its peculiar metapopulation structure and the peculiar mixture and strength of homeostatic processes vis à vis heterostatic ones—that determines species’ identity at a time and through time. We then explore the consequences of the acceptance of the empirical approach in settling the individuals versus kinds dispute. In particular, (...) while conceptual arguments have been proposed to show that species can be equally treated as individuals and kinds because mereology’s and set-theory’s languages are inter-translatable, we advance instead a causal argument to sustain the claim that each species is both a kind and an individual. (shrink)

In this paper, I identify two general positions with respect to the relationship between environment and natural selection. These positions consist in claiming that selective claims need and, respectively, need not be relativized to homogenous environments. I then show that adopting one or the other position makes a difference with respect to the way in which the effects of selection are to be measured in certain cases in which the focal population is distributed over heterogeneous environments. Moreover, I show that (...) these two positions lead to two different interpretations—the Pricean and contextualist ones—of a type of selection scenarios in which multiple groups varying in properties affect the change in the metapopulation mean of individual-level traits. Showing that these two interpretations stem from different attitudes towards environmental homogeneity allows me to argue: that, unlike the Pricean interpretation, the contextualist interpretation can only claim that drift or selection is responsible for the change in frequency of the focal trait in a given metapopulation if details about whether or not group formation is random are specified; that the traditional main objection against the Pricean interpretation—consisting in arguing that the latter takes certain side-effects of individual selection to be effects of group selection—is unconvincing. This leads me to suggest that the ongoing debate about which of the two interpretations is preferable should concentrate on different issues than previously thought. (shrink)

The concept of population is central to ecology, yet it has received little attention from philosophers of ecology. Furthermore, the work that has been done often recycles ideas that have been developed for evolutionary biology. We argue that ecological populations and evolutionary populations, though intimately related, are distinct, and that the distinction matters to practicing ecologists. We offer a definition of ecological population in terms of demographic independence, where changes in abundance are a function of birth and death processes alone. (...) However, demographic independence is insufficient on its own so we supplement it with the idea of shared habitat. An ecological population is a group of organisms of the same species in a habitat that manifests DI. Given the importance of metapopulation dynamics to modern ecology, an account of ecological population must apply to this domain as well. Thus, we extend our definition of ecological population to the metapopulation. To facilitate the extension, we introduce the metahabitat—a collection of spatially segregated habitat patches shared by a single DI population. This enables us to diagnose some unhelpful trends in the metapopulation literature and emphasize the importance of habitat dynamics in pursuit of the goals of theoretical ecology and conservation biology. (shrink)

The issue of whether formal reasoning or a computing-intensive approach is the most efficient manner to address scientific questions is the subject of some considerable debate and pertains not only to the nature of the phenomena and processes investigated by scientists, but also the nature of the equation and algorithm objects they use. Although algorithms and equations both rely on a common background of mathematical language and logic, they nevertheless possess some critical differences. They do not refer to the same (...) level of symbolization, as equations are based on integrated concepts in a denotational manner, while algorithms specifically break down a complex problem into more elementary operations, in an operational manner. They may therefore be considered as suited to the representation of different phenomena. Specifically, algorithms are by nature sufficient to represent weak emergent phenomena, but not strong emergent patterns, while equations can do both. Finally, the choice between equations and algorithms are by nature sufficient to represent weak emergent phenomena, but not strong emergent patterns, while equations behave conversely. We propose a simplified classification of scientific issues for which both equation- and/or algorithm-based approaches can be envisaged, and discuss their respective pros and cons. We further discuss the complementary and sometimes conflicting uses of equations and algorithms in a context of ecological theory of metapopulation dynamics. We finally propose both conceptual and practical guidelines for choosing between the alternative approaches. (shrink)

At least three different issues are commonly referred to by the term “the species problem”: one concerns the necessary properties of species, a second the processes responsible for the existence of species, and a third methods for inferring species limits. Solutions have recently been proposed to the first two problems, which are conceptual in nature (the third is methodological). The first equates species with metapopulation lineages and proposes that existence as a separately evolving metapopulation lineage be considered the only necessary (...) property of species. The second views the species category as a cluster concept and proposes that no single process or set of processes be considered necessary for the existence of species. Although these two solutions have been portrayed as being in conflict, they are, in fact, highly compatible. Moreover, the proposals in question clarify the problem concerning methods for inferring the limits of species, which has for a long time been confused with the problem concerning the necessary properties of species. Together these proposals provide the opportunity for biology to move beyond debates about the definition of the species category and focus on estimating the boundaries and numbers of species as well as studying the diverse processes involved in their origin and persistence. BioEssays 27:1263–1269, 2005. © 2005 Wiley Periodicals, Inc. (shrink)

The novel coronavirus pandemic is intensifying all over the world, but some countries, including China, have developed extensive and successful experience in controlling this pandemic. In this context, some questions arise naturally: What can countries caught up in the epidemic learn from China’s experience? In regions where the outbreak is under control, what would lead to a resurgence of the epidemic? To address these issues, we investigate China’s experience in anticontagion interventions and reopening process, focusing on the coevolution of epidemic (...) and awareness during COVID-19 outbreak. Through an empirical analysis based on large-scale data and simulation based on a metapopulation and multilayer network model, we ascertain the impact of human movements and awareness diffusion on the epidemic, elucidate the inherent patterns and effective interventions of different epidemic prevention methods, and highlight the crunch time of each measure. The results are also employed to analyze COVID-19 evolution in other countries so as to find unified rules in complex situations around the world and provide advice on anticontagion and reopening policies. Our findings explain some key mechanisms of epidemic prevention and may help the epidemic analysis and decision-making in various countries suffering from COVID-19. (shrink)

The variance effective population size (NeV\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$N_{eV}$$\end{document}) is frequently used to quantify the expected rate at which a population’s allele frequencies change over time. The purpose of this paper is to find expressions for the global NeV\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$N_{eV}$$\end{document} of a spatially structured population that are of interest for conservation of species. Since NeV\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$N_{eV}$$\end{document} depends on allele (...) frequency change, we start by dividing the cause of allele frequency change into genetic drift within subpopulations (I) and a second component mainly due to migration between subpopulations (II). We investigate in detail how these two components depend on the way in which subpopulations are weighted as well as their dependence on parameters of the model such a migration rates, and local effective and census sizes. It is shown that under certain conditions the impact of II is eliminated, and NeV\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$N_{eV}$$\end{document} of the metapopulation is maximized, when subpopulations are weighted proportionally to their long term reproductive contributions. This maximal NeV\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$N_{eV}$$\end{document} is the sought for global effective size, since it approximates the gene diversity effective size NeGD\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$N_{eGD}$$\end{document}, a quantifier of the rate of loss of genetic diversity that is relevant for conservation of species and populations. We also propose two novel versions of NeV\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$N_{eV}$$\end{document}, one of which (the backward version of NeV\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$N_{eV}$$\end{document}) is most stable, exists for most populations, and is closer to NeGD\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$N_{eGD}$$\end{document} than the classical notion of NeV\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$N_{eV}$$\end{document}. Expressions for the optimal length of the time interval for measuring genetic change are developed, that make it possible to estimate any version of NeV\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$N_{eV}$$\end{document} with maximal accuracy. (shrink)

Recently the role of customary local institutions in the conservation of biological diversity has become a topic of widespread interest. In this paper the conservation value of one such institution, traditionally protected forest, is studied with regard to its ecological representativity and institutional persistence. On the basis of a case study from Mozambique the paper concludes that traditionally protected forests do have a practical conservation value, especially as fire refuges and in the preservation of metapopulations of endangered species. However, (...) it is also important to recall their spiritual aspect, which is crucial for their continuing appreciation and upholding. Even though customary institutions are still strong in the study area, they are subject to power struggles at both local and national levels. In the present context of political transition, any outside interventions regarding such local institutions as sacred forests, which have high symbolic value, should be considered carefully. (shrink)

This afterword to Species and Beyond provides some reflections on species, with special attention to what I think the most significant developments have been in the thinking of biologists and philosophers working on species over the past 25 years, as well as some bad jokes.

Ernst Mayr played a central role in the establishment of the general concept of species as metapopulation lineages, and he is the author of one of the most popular of the numerous alternative definitions of the species category. Reconciliation of incompatible species definitions and the development of a unified species concept require rejecting the interpretation of various contingent properties of metapopulation lineages, including intrinsic reproductive isolation in Mayr's definition, as necessary properties of species. On the other hand, the general concept (...) of species as metapopulation lineages advocated by Mayr forms the foundation of this reconciliation, which follows from a corollary of that concept also advocated by Mayr: the proposition that the species is a fundamental category of biological organization. Although the general metapopulation lineage species concept and Mayr's popular species definition are commonly confused under the name "the biological species concept," they are more or less clearly distinguished in Mayr's early writings on the subject. Virtually all modern concepts and definitions of the species category, not only those that require intrinsic reproductive isolation, are to be considered biological according to the criterion proposed by Mayr. Definitions of the species category that identify a particular contingent property of metapopulation lineages (including intrinsic reproductive isolation) as a necessary property of species reduce the number of metapopulation lineages that are to be recognized taxonomically as species, but they cause conflicts among alternative species definitions and compromise the status of the species as a basic category of biological organization. (shrink)

The scenario of Homo sapiens origin/s within Africa has become increasingly complex, with a pan-African perspective currently challenging the long-established single-origin hypothesis. In this paper, we review the lines of evidence employed in support of each model, highlighting inferential limitations and possible terminological misunderstandings. We argue that the metapopulation scenario envisaged by pan-African proponents well describes a mosaic diversification among late Middle Pleistocene groups. However, this does not rule out a major contribution that emerged from a single population where crucial (...) derived features—notably, a globular braincase—appeared as the result of a punctuated, cladogenetic event. Thus, we suggest that a synthesis is possible and propose a scenario that, in our view, better reconciles with consolidated expectations in evolutionary theory. These indicate cladogenesis in allopatry as an ordinary pattern for the origin of a new species, particularly during phases of marked climatic and environmental instability. (shrink)

Ontological understanding of biological units is crucial to their use in experimental design, analysis, and interpretation. Conceptualizing fundamental units in biology as individuals or classes is important for subsequent development of discovery operations. While the criteria for diagnosing individuals are acknowledged, temporal boundedness is often misinterpreted and temporal minima are applied to units in question. This results in misdiagnosis or abandonment of ontological interpretation altogether. Biological units such as areas of endemism in biogeography and species in evolutionary biology fall victim (...) to such problems. Our goal here is to address the misconception that biological individuals such as species and areas of endemism have a temporal minimum. Areas of endemism can persist within small temporal boundaries in the context of metapopulation dynamics, island biogeography, and range expansion and contraction. Similarly, lineage reticulation illustrates examples of short-lived species. Here, examples of known entities are provided to illustrate their persistence on short time scales in attempt to rescue future interpretation of biological units from ontological misdiagnosis, elucidate the philosophical individuality of areas of endemism and species with short lifespans, and provide justification for the “snapshot in time” diagnostic approach. (shrink)

This paper illuminates a variety of issues that speak to the question of whether ‘captivity for conservation’ can be an ethically acceptable goal of the modern zoo. Reflecting on both theoretical disagreements and practical challenges , the paper explains why the ‘Noah’s Ark’ paradigm is being replaced by an alternative ‘integrated approach.’ It explores the changes in the zoo’s core tasks that the new paradigm implies. And it pays special attention to the changes that would have to be made in (...) zoos’ collection policies: connection with in situ projects, emphasizing local species and the local biogeographical region, exchange of animals among zoos and between zoos and wildlife, and a shift towards smaller species. Finally the question will be addressed whether the new paradigm will achieve a morally acceptable balance between animal welfare costs and species conservation benefits. (shrink)