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Ecotone Formation Through Ecological Niche Construction: the Role of Biodiversity and Species Interactions. Kévin Liautaud, Matthieu Barbier, Michel Loreau

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Ecotone Formation Through Ecological Niche Construction: the Role of Biodiversity and Species Interactions. Kévin Liautaud, Matthieu Barbier, Michel Loreau Ecotone formation through ecological niche construction: the role of biodiversity and species interactions. Kévin Liautaud, Matthieu Barbier, Michel Loreau To cite this version: Kévin Liautaud, Matthieu Barbier, Michel Loreau. Ecotone formation through ecological niche con- struction: the role of biodiversity and species interactions.. Ecography, Wiley, 2020, 45 (5), pp.714- 723. 10.1111/ecog.04902. hal-02491373 HAL Id: hal-02491373 https://hal.archives-ouvertes.fr/hal-02491373 Submitted on 26 Feb 2020 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. 43 1–10 ECOGRAPHY Research Ecotone formation through ecological niche construction: the role of biodiversity and species interactions Kevin Liautaud, Matthieu Barbier and Michel Loreau K. Liautaud (https://orcid.org/0000-0001-6164-8415) ✉ ([email protected]), M. Barbier and M. Loreau, Centre for Biodiversity Theory and Modelling, Theoretical and Experimental Ecology Station, UMR 5321, CNRS and Paul Sabatier Univ., Moulis, France. Ecography Rapid changes in species composition, also known as ecotones, can result from vari- 43: 1–10, 2020 ous causes including rapid changes in environmental conditions, or physiological doi: 10.1111/ecog.04902 thresholds. The possibility that ecotones arise from ecological niche construction by ecosystem engineers has received little attention. In this study, we investigate how Subject Editor: Dominique Gravel the diversity of ecosystem engineers, and their interactions, can give rise to ecotones. Editor-in-Chief: We build a spatially explicit dynamical model that couples a multispecies community Jens-Christian Svenning and its abiotic environment. We use numerical simulations and analytical techniques Accepted 12 January 2020 to determine the biotic and abiotic conditions under which ecotone emergence is expected to occur, and the role of biodiversity therein. We show that the diversity of ecosystem engineers can lead to indirect interactions through the modification of their shared environment. These interactions, which can be either competitive or mutual- istic, can lead to the emergence of discrete communities in space, separated by sharp ecotones where a high species turnover is observed. Considering biodiversity is thus critical when studying the influence of species–environment interactions on the emer- gence of ecotones. This is especially true for the wide range of species that have small to moderate effects on their environment. Our work highlights new mechanisms by which biodiversity loss could cause significant changes in spatial community patterns in changing environments. Keywords: biodiversity, community patterns, ecological niche construction, ecosystem engineers, ecotones, species interactions Introduction Whether species composition changes gradually, or forms discrete zones along environ- mental gradients has been the subject of a long-standing debate in ecology (Clements 1916, Gleason 1926, Braun-Blanquet 1928, Hedberg 1955, McIntosh 1967). Observational studies have found both gradual (Whittaker 1956, Lieberman et al. 1996, Vazquez and Givnish 1998, Ellison et al. 2010) and discrete patterns (Kitayama 1992, Tuomisto and Ruokolainen 1994, Kessler 2000, Hemp 2006). Rapid changes in community composition along gradients, also termed ecotones (Kent et al. 1997), have been observed in a wide range of ecosystems, such as alpine treelines (Germino et al. 2002), tropical mountain forests (Martin et al. 2007) and coastal environments –––––––––––––––––––––––––––––––––––––––– © 2020 The Authors. Ecography published by John Wiley & Sons Ltd on behalf of Nordic Society Oikos www.ecography.org This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. 1 (Walker et al. 2003, Sternberg et al. 2007). Hereafter, a tran- Biodiversity can have two main effects on the emergence sition will be termed ‘rapid’ when its scale is much smaller of species–environment feedbacks: a cumulative effect of than the spatial scale of the landscape, even though the tran- species number, and a heterogeneity effect due to variations sitional area may show mixing of species. in species’ preferences and engineering ability. Cumulative While rapid changes can be blurred by species dispersal effects are similar to complementarity in biodiversity–eco- (Liautaud et al. 2019) or stochasticity in nature, it is impor- system functioning relationships (Loreau and Hector 2001, tant to understand the theoretical conditions under which Hooper et al. 2005). The fact that species coexist with weak or rapid community changes can emerge. These rapid changes no competition implies the existence of different niches, i.e. in species composition can coincide with rapid changes in other factors beyond the environmental preference modelled environmental conditions, such as the frost line (Kitayama here. This cumulative effect is most important when there is and Mueller-Dombois 1992) or discontinuities in edaphic no single identifiable engineer, but where a community acts conditions (Tuomisto and Ruokolainen 1994, Kessler 2000). collectively to create an ecotone. A potential example is the In these cases, it is often assumed that changes in abiotic occurrence of ecotones between mangroves and hardwood conditions are responsible for the change in species composi- forests, where several mangrove tree species can modify water tion (McIntosh 1967, Kent et al. 1997). This assumption is salinity in synergy (Sternberg et al. 2007). In contrast, the supported in many cases, but it may obscure the possibility heterogeneity effect of biodiversity arises when there are dif- that, in other settings, the two boundaries emerge together ferences in species’ preferred environmental states. We inves- from the influence of species on their abiotic environment. tigate the effect of these differences on emergent competition The mechanisms that can lead to such transitions are poorly or facilitation between ecosystem engineers, and how this known, and in particular the respective contributions of spe- could play a role in ecotone emergence. cies–environment feedbacks and interspecific interactions. In this study, we build a theoretical model that couples the Species that are able to modify their abiotic environment dynamics of a community and of its abiotic environment to are often called ‘ecosystem engineers’ (Jones et al. 2010). assess the role of ecosystem engineers and of their diversity in Classical examples range from beavers that impact water flow the emergence of ecotones in space. In our model, ecotones and habitat heterogeneity (Wright et al. 2002), to cushion are represented by abrupt changes, including discontinui- alpine plants that buffer extreme temperatures and increase ties. In the presence of multiple interacting species, we show soil moisture (Badano et al. 2006). Ecological niche con- that ecological niche construction can lead to the emergence struction is a particular case in which engineers modify of indirect interspecific interactions – which can be either the environment to their own benefits (Kylafis and Loreau positive or negative – through environmental modifications. 2008, 2011), creating a feedback with the environment (an Similarly, we show that even species with different prefer- example in which engineers can instead create succession is ences can act synergistically as a single community. We then presented in the Supplementary material Appendix 1–5). assess the consequences of these different interaction types This ecological process should be distinguished from the for community patterns in space, and identify the conditions related concept of niche construction in evolutionary theory under which ecotone formation is predicted to occur. in which we would also expect species traits to evolve over time (Odling-Smee et al. 1996, 2003). Examples of ecologi- cal niche construction range from plant–water feedbacks in Model and methods arid environment (Dekker et al. 2007) to increases in nutri- ent inputs by trees in tropical ecosystems (De longe et al. Species growth and niche construction 2008). Such feedbacks can govern species distributions (Wilson and Agnew 1992), particularly under harsh environ- We model the dynamics of a community of n species, each mental conditions (von Hardenberg et al. 2001, Gilad et al. of which obeys a logistic growth along a gradient of an arbi- 2004, Meron et al. 2004, Kéfi et al. 2007), and lead to the trary environmental factor E. We consider independent loca- emergence of ecotones (Jiang and DeAngelis 2013, Bearup tions along this environmental gradient, assuming no fluxes and Blasius 2017). Classical studies on ecosystem engineers, between the locations (but see Liautaud et al. (2019) for the however, have generally focused on the effects of a particu- role of dispersal in smoothing abrupt transitions). For a given lar species having strong effects on the abiotic environment location k, the population dynamics
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