Community Stability and Turnover in Changing Environments Kevin Liautaud

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Community Stability and Turnover in Changing Environments Kevin Liautaud Community stability and turnover in changing environments Kevin Liautaud To cite this version: Kevin Liautaud. Community stability and turnover in changing environments. Biodiversity and Ecology. Université Paul Sabatier - Toulouse III, 2020. English. NNT : 2020TOU30264. tel- 03236079 HAL Id: tel-03236079 https://tel.archives-ouvertes.fr/tel-03236079 Submitted on 26 May 2021 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. THTHESEESE`` En vue de l’obtention du DOCTORAT DE L’UNIVERSITE´ DE TOULOUSE D´elivr´epar : l’Universit´eToulouse 3 Paul Sabatier (UT3 Paul Sabatier) Pr´esent´eeet soutenue le 13 Mars 2020 par : Kevin LIAUTAUD Community stability and turnover in changing environments JURY THEBAUD Christophe Professeur d’Universit´e Pr´esident du Jury BARABAS Gyorgy¨ Professeur Assistant Membre du Jury POGGIALE Jean-Christophe Professeur d’Universit´e Membre du Jury THEBAULT Elisa Charg´eede Recherche Membre du Jury SCHEFFER Marten Directeur de Recherche Membre du Jury LOREAU Michel Directeur de Recherche Membre du Jury Ecole´ doctorale et sp´ecialit´e: SEVAB : Ecologie,´ biodiversit´eet ´evolution Unit´ede Recherche : CNRS-Universit´ePaul Sabatier, UMR 5321, Station d’Ecologie Th´eorique& Exp´erimentale,2 Route du CNRS, 09200 Moulis, France Directeur(s) de Th`ese: Michel Loreau et Marten Scheffer Rapporteurs : Jean-Christophe Poggiale et Gy¨orgy Barab´as 2 Cette th`esea ´et´er´ealis´eeau sein du laboratoire: Station d’Ecologie Th´eorique et Exp´erimentale (SETE) du CNRS (UMR 5321, CNRS, Universit´ePaul Sabatier) 2, route du CNRS, 09200 Moulis (France) Elle a ´et´efinanc´eepar le projet ERC BIOSTASES : BIOdiversity, STAbility and sustainability in Spatial Ecological and social-ecological Systems (Num´ero: 666971 ), et le Laboratoire d’Excellence TULIP (Towards a unified theory of biotic interactions). 3 Abstract The question whether communities should be viewed as superorganisms or loose collections of individual species has been the subject of a long-standing debate in ecology. Each view implies different spatial and temporal community patterns. When environment gradually changes in space or in time, the organis- mic view predicts that species turnover is discontinuous, while the individualis- tic view predicts gradual changes in species composition. The main objective of this thesis is to understand the theoretical conditions under which these various types of community response can occur. First, I study the role of interspecific competition can play in the emergence of various spatial community patterns. I investigate the theoretical conditions in competition under which smooth or discrete spatial patterns can emerge. Then, I study how interactions between species and their environment can lead to various community patterns in space. I notably show how ecological niche construction can lead to the emergence of abrupt changes in species composition and in the environment, and the role bio- diversity plays therein. Finally, I focus on the role biodiversity can play against ecosystem collapse. In this section, I illustrate how diversity loss, through its effects on total biomass, can lead to ecosystem collapse. 4 R´esum´e Les communaut´esd’ˆetresvivants peuvent elles ˆetres consid´er´eescomme des organismes complexes, ou au contraire comme de simples groupes d’esp`eces,in- dividuelles ? Cette question est `al’origine de nombreux d´ebatsen ´ecologie, ces deux visions impliquant notamment des pr´edictionstr`esdiff´erentes dans les patrons spatiaux et temporels de communaut´es. Lorsque l’environnement change graduellement dans l’espace ou dans le temps, la vision individualiste im- plique des changements graduels dans la composition des communaut´es,tandis que la vision du ”super-organisme” pr´editdes changements davantage abrupts. L’objectif principal de cette th`eseest de comprendre et d´eterminersous quelles conditions ces diff´erents types de r´eponse des communaut´es aux changements de l’environnement peuvent advenir. Dans une premi`erepartie, nous ´etudionsle rˆoleque la comp´etitioninter-sp´ecifiquepeut jouer dans l’´emergencede diff´erents patrons spatiaux de communaut´es. Nous ´etudionsnotamment les conditions th´eoriquessous lesquelles la comp´etitionpeut faire apparaitre des patrons gradu- els ou discontinus dans la composition des esp`eces. Dans une deuxi`emepartie, nous ´etudionsl’influence des interactions entre les esp`eceset leur environnement sur les patrons spatiaux de communaut´es.Nous montrons notamment comment des ph´enom`enesde construction de niche peuvent mener `al’´emergencede change- ments brutaux dans la composition des communaut´es,mais ´egalement dans les conditions de l’environnement. Enfin, dans une derni`erepartie, nous illustrons le rˆoleque peut jouer la biodiversit´edans la protection des ´ecosyst`emesface `a des effondrements ´ecologique,et notamment le rˆoleque peut jouer la biomasse dans cette protection. 5 Remerciements Peu de choses sont aussi collectives qu’un doctorat en sciences. Je souhaiterai remercier ici toutes les personnes qui ont rendu possible l’aboutissement de ce projet de trois ann´ees. Je tiens premi`erement `aremercier Michel, mon directeur de th`ese,de m’avoir accord´esa confiance dans la r´ealisationde ce projet. Merci pour la libert´eque tu m’as laiss´edans la conduite de ce travail, et pour les conseils avis´esdont j’ai pu b´en´eficierpendant ces trois ann´eesau sein de la SETE. I also would like to thanks Marten Scheffer, for having accepted to be co-director on this project. Thank you for having supervised and helped me and for your invitations to work with your team in Wageningen. I also would like to thank Egbert van Nes for his help during our scientific collaborations, and for having been part of my PhD Comitee. Je tiens ´egalement `aremercier Wilfried Thuiller et Sonia K´efipour leurs conseils lors des comit´esde suivi de cette th`ese. Mes remerciements vont ´egalement `a Jean-Christophe Poggiale et Gy¨orgyBarab´as,qui ont accept´ed’ˆetrerapporteurs de ce travail, ainsi qu’`aElisa Thebault et Christophe Thebaud, membres du Jury. Mes remerciements vont ´egalement `al’Europen Research Council, qui a financ´ecette th`esede doctorat au sein du projet BIOSTASES, et m’a permis de la r´ealiserdans les meilleures conditions. Parmi les personnes qui ont contribu´ela r´ealisationde cette th`ese,je souhait- erai remercier Dalila Booth pour avoir grandement facilit´ela gestion adminis- trative de ce projet, ainsi que Marion Bousquet, Myriam Istoczak et S´everine Bonzom. Je tiens ´egalement `aremercier le CNRS et l’ensemble de ses person- nels. Mes remerciements vont ´egalement `aDominique Pantalacci, de l’Ecole Doctorale SEVAB, qui a grandement facilit´eles d´emarches administratives et li´eesaux formations. J’ai eu la chance de b´en´eficierde l’aide et de l’inspiration de nombreuses 6 personnes au sein de cette station. Je tiens `aremercier particuli`erement Jeff et Yuval pour nos dicussions, et pour leurs ´eclairagessur des questions scientifiques qui me d´epassaient souvent. Tous mes remerciements vont ´egalement `aMatthieu, qui m’a tr`esgrandement aid´eet inspir´edurant l’ensemble de ces trois ans, et dont la culture scientifique et l’enthousiasme ont ´et´eabsolument d´ecisifsdans la r´eussitede ce projet, de la premi`ereid´eejusqu’`ala publication. Les id´eesviennent parfois l`ao`uon les attend le moins, et beaucoup d’entre elles sont venues en compagnie des nombreuses personnes avec qui j’ai pu partager un caf´e,une sortie en montagne ou naturaliste. Merci `ala team muffilm - caf´e- wrap du march´e: Pierre, Nuria, Kirsten, David, Vini, Laurane, Rik, Lea, Math- ieu, Maya, Jo, Yuval, Matthieu, Anne-So, K´eoni,J´erˆome,Aisha, Robin, Hugo, Elo et Nico. Merci `aJo, Jeff, Nuria, K´eoni,Diego, Pauline, Th´eo,Az et Anne-So pour les ours cach´es,le calcaire tremp´e, les arˆetesen granite et le remonte- pente du Riberot. Mes remerciements vont ´egalement `aStefano et Rocco, qui ont su m’inspirer quand je d´ebutaisla recherche dans les Alpes italiennes, ainsi qu’`aSylvie et Hendrik avec qui j’ai d´ebut´ela mod´elisationen ´ecologie. Merci ´egalement aux amis de longue date dont les retours au pays comme les sorties en altitude ont toujours constitu´esdes pauses privil´egi´ees et salvatrices. Lucien, Julie, PA, Lor`ene,Val, Thomas J, TG, C´esar,R´emi,et la grande et belle ´equipe d’Auzeville, Grazie per tutto ! Mille ”merci” reconnaissants `aMarie, qui a su me donner une incroyable ´energie lorsque le doute et le d´ecouragement prenaient le dessus. Ce projet n’aurait pas pu aboutir sans tes encouragements et ton soutien ind´efectible. Merci pour tout. Merci enfin `ames parents, `aMarine, et `al’ensemble de ma famille pour leur soutien sans failles durant ces ann´eesd’´etudeet de travail, quelquefois sem´eesd’embˆuches, et de choix d´ecisifs.Vous avez su m’aider `afaire les bons. Merci beaucoup. KL 7 8 Contents Introduction g´en´erale 11 General Introduction 29 1 Superorganisms or loose collections of species? A unifying the- ory of community patterns along environmental gradients 44 1.1 Abstract . 45 1.2 Introduction . 46 1.3 Model and Methods . 49 1.3.1 Competition . 49 1.3.2 Dispersal . 50 1.3.3 Species turnover and changes in species abundances . 51 1.3.4 Effects of competition on community properties and com- munity attractors . 52 1.3.5 Numerical simulations . 53 1.4 Results . 54 1.4.1 Effects of competition on spatial community patterns with- out dispersal . 54 1.4.2 Effects of dispersal on spatial community patterns .
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