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How Individualized Niches Arise: Defining Mechanisms of Niche Construction, Niche Choice and Niche Conformance

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How Individualized Niches Arise: Defining Mechanisms of Niche Construction, Niche Choice and Niche Conformance 1How Individualized Niches Arise: Defining Mechanisms of Niche 2Construction, Niche Choice and Niche Conformance 3Rose Trappes*, Behzad Nematipour*, Marie I. Kaiser*, Ulrich Krohs*, Koen J. van 4Benthem^, Ulrich R. Ernst^, Jürgen Gadau^, Peter Korsten^, Joachim Kurtz^, Holger 5Schielzeth^, Tim Schmoll^, Elina Takola^ 6 7*Shared first authorship 8^Equal contribution 9 10Affiliations 11RT, MK: Department of Philosophy, Bielefeld University, Universitätsstr. 25, 33615 12Bielefeld, Germany 13BN: Center for Philosophy of Science, University of Münster, Domplatz 23, 48143 Münster, 14Germany 15UK: Department of Philosophy, University of Münster, Domplatz 23, 48143 Münster, 16Germany 17KvB: Department of Theoretical Biology, Bielefeld University, Universitätsstr. 25, 33615 18Bielefeld, Germany 19UE, JG, JK: Institute for Evolution and Biodiversity, University of Münster, Hüfferstraße 1, 2048149 Münster, Germany 21PK: Department of Animal Behaviour, Bielefeld University, Konsequenz 45, 33615 22Bielefeld, Germany 23TS: Department of Evolutionary Biology, Bielefeld University, Konsequenz 45, 33615 24Bielefeld, Germany 25HS, ET: Population Ecology Group, Institute of Ecology and Evolution, Friedrich Schiller 26University Jena, Dornburger Straße 159, 07743 Jena, Germany 27 28ORCiD 29RT: 0000-0002-6398-5404 30BN: - 31MIK: 0000-0002-9307-7474 32UK: - 33KvB: 0000-0002-3841-2110 34UE: 0000-0002-6330-5341 35JG: 0000-0003-1258-5379 36PK: 0000-0003-0814-9099 37JK: 0000-0002-7258-459X 1 38HS: 0000-0002-9124-2261 39TS: 0000-0003-3234-7335 40ET: 0000-0003-1268-5513 41 42Correspondence 43Rose Trappes, [email protected] 44 45Author Contribution Statement 46All authors conceived the theoretical framework. RT, BN, MK & UK created the initial draft 47and performed revisions of the manuscript. KvB, UE, JG, PK, JK, HS, TS & ET contributed 48examples and helped to revise the text and figures. All authors approved the submitted 49version. 50 51Funding 52This research was funded by the German Research Foundation (DFG) as part of the SFB 53TRR 212 (NC³) – Project number 316099922. 54 55Acknowledgements 56We would like to acknowledge Caroline Müller, Oliver Krüger, and Norbert Sachser for 57generating the initial framework of the NC3 mechanisms which we develop and present here, 58and for helpful comments during writing. Thanks are also due to all members of the CRC 59TRR-212 at the University of Münster, Bielefeld University and Jena University and to the 60philosophers of biology at Bielefeld University for their attention and constructive feedback 61on the project. 2 62How Individualized Niches Arise: Defining Mechanisms of Niche Construction, Niche 63Choice and Niche Conformance 64 65Abstract 66The debate between the extended evolutionary synthesis (EES) and the modern synthesis 67(MS) partly relies on different interpretations of niche construction. We dissect the umbrella 68term of niche construction into three separate mechanisms: niche construction (taken in a 69narrow sense), in which individuals make changes to the environment; niche choice, in which 70individuals select an environment; and niche conformance, in which individuals change their 71phenotypes. Each of these individual-level mechanisms affects an individual’s phenotype- 72environment match, its fitness, and its individualized niche, defined in terms of the 73environmental conditions under which an individual can survive and reproduce. Our 74conceptual framework distinguishes several ways in which individuals alter the selective 75regimes that they and other organisms experience. It also places clear emphasis on individual 76differences and construes niche construction and other processes as evolved mechanisms. We 77therefore argue that our framework helps to resolve the tensions between EES and MS. 78 79Keywords: extended evolutionary synthesis, niche construction, individualized niche, 80individual differences, phenotypic plasticity 81 82Introduction 83Organisms change their environment—beavers build dams, birds build nests, earthworms 84alter soil structure. It seems uncontentious that these organisms thereby modify their living 85conditions and the selective regime under which they live. Under dispute, however, is what 86these modificatory actions, often called niche construction, mean for evolutionary theory. 3 87One view is that through modifying the environment organisms actively influence the 88evolution of their own and other species, making organisms not just objects of, but subjects in 89evolution (Lewontin 1983, Laland et al. 2015). This view is central to the Extended 90Evolutionary Synthesis (EES). Another view interprets organisms’ modificatory actions as 91adaptations that are the outcome of earlier stages of evolution. This classical view of the 92Modern Synthesis (MS), in contrast to EES, therefore maintains organisms’ status as mere 93objects of evolutionary processes, with environmental modifications understood as a result of 94evolution by natural selection (Dawkins 1982). 95The dispute might look like a chicken and egg problem—which came first, selection or 96construction? But there is far more at stake than just where to put the focus in a sequence of 97alternating steps. EES and MS have starkly contrasting understandings of evolution and take 98different biological disciplines as relevant for evolutionary theory. EES considers organismal 99activities as dependent on modifications in developmental processes, which in turn affect 100evolution (Laland et al. 2014). It thus demands an extension of MS by, for instance, 101integrating developmental and behavioral biology. In contrast, contemporary interpretations 102of MS acknowledge developmental processes and interactions between the genotype and the 103environment, yet claim that these can be accommodated within the classical framework that 104sees genetic change as the core of evolutionary change (Wray et al. 2014). 105We believe that the alleged incompatibility of EES and MS accounts of niche construction 106can be partly resolved. We argue for recognizing the different ways in which organisms alter 107their environment and their relation to the environment as a set of evolved causal mechanisms 108that play a role in evolutionary processes. This insight derives from our interdisciplinary 109work within the Research Consortium “A Novel Synthesis of Individualisation across 110Behaviour, Ecology and Evolution: Niche Choice, Niche Conformance and Niche 111Construction (NC3)” (https://www.uni-bielefeld.de/biologie/crc212). In this paper, we 4 112undertake some conceptual clarifications that help to diminish the perceived conflict between 113EES and MS. We do so largely with a focus on animals, but we see our framework as 114applicable to plants and other taxa. 115We first point out that niche construction is carried out by individuals and that there can be 116individual differences in niche construction (section 1). We subdivide the umbrella concept 117of niche construction into three distinct mechanisms through which individuals interact with 118their environments: making changes to the environment (niche construction in the narrow 119sense), selecting an environment (niche choice), and altering their phenotype in response to 120the environment (niche conformance) (sections 2 and 3). All three mechanisms alter the 121phenotype-environment match and the individual’s fitness, thereby determining the 122ecological niche of an individual, defined as the set of environmental conditions in which an 123individual can survive and reproduce (section 4). Hence, we refer to these mechanisms as 124“niche-altering mechanisms” or “NC3 mechanisms”. Finally, we discuss which aspects of the 125debate about the evolutionary relevance of niche construction can be resolved by our 126conceptual work, which ones become empirical questions, and which ones remain as 127differences between different framings of evolution (section 5). 128 1291. Individual Differences in Niche Construction 130We can start with a classical example of niche construction: “Birds build nests” or more 131concretely, “Common buzzards (Buteo buteo) build nests with twigs from various tree 132species.” Here, niche construction might be understood at the species level, describing what 133nest-building behavior is characteristic of the species B. buteo. Yet in the end it is individual 134buzzards that build nests in the environment. In fact, niche construction activities always 135require the action of individuals (sometimes in pairs or groups) that change the environment. 5 136According to the EES, these individual activities can in turn affect the evolutionary 137trajectories of populations and species. 138The relation between individual activities and species evolution is sometimes obscured in 139definitions of niche construction. For instance, Lewontin and Levins first described niche 140construction—prior to the coining of the term—as such: “the organism influences its own 141evolution, by being both the object of natural selection and the creator of the conditions of 142that selection” (Lewontin and Levins 1985, p. 106). Here “the organism” is at once 143population or species—since it has its “own” evolution, and it is populations and species 144which evolve—and individual—performing activities that alter the conditions of selection. 145Later definitions of niche construction try to avoid this equivocation. For instance, Odling- 146Smee, Laland and Feldman caution that “niche construction is typically expressed by 147individual organisms, but natural
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