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Theoretical and Experimental Ecology Station

TheoMoDive

TheoMoDive

Biodiversity Theory and Modeling Research Group

Presentation

The erosion of biodiversity is, along with climate change, one of the greatest challenges societies will face in this century and beyond (Dirzo et al. 2014, Hallmann et al. 2017, Ripple et al. al. 2017, IPBES 2019, IPCC 2019). Indeed, biodiversity is part of most human activities, directly or indirectly, by providing biological resources used for multiple purposes (food and health being the most obvious examples), by helping to maintain a wide range of ecological services on which all human activities depend, and by participating in the very framework of the material and spiritual life of the human species. The ongoing erosion of biodiversity is therefore likely to have far-reaching consequences for societies in the centuries to come. It is in this context that an integrative science of biodiversity is emerging (Loreau 2010), whose links with decision-making processes should be strengthened, in particular through the Intergovernmental Platform on Biodiversity and Services. Ecosystems (IPBES).
Theoretical ecology plays an essential role in shedding light on the cause and effect processes and relationships that take place in the particularly complex systems of ecosystems and socio-ecological systems. It also helps to better understand the impacts of changes in biodiversity by clarifying complex concepts, fundamental to the study of biodiversity and its causes and consequences, such as biodiversity, functioning and stability of ecosystems.
France has a strong community of theoretician and modelling ecologists, and the meetings of the GDR Theory and Modeling of Biodiversity play an important unifying role in this community. Our community finds considerable interest in comparing the approaches and coordinating the research efforts of the various research teams in France in this field.

Research Themes

ThEme 1. DIVERSITY, STABILITY AND FUNCTIONING OF ECOLOGICAL SYSTEMS

ThEme 2. DYNAMICS OF INTERACTION NETWORKS

ThEme 3. SPATIAL DYNAMICS, METACOMMUNITIES AND META-ECOSYSTEMS

THEME 4. DYNAMICS OF INTERACTIONS BETWEEN BIODIVERSITY AND SOCIETY

THEME 5. THE ROLE OF EVOLUTION

THEME 1. DIVERSITY, STABILITY AND FUNCTIONING OF ECOLOGICAL SYSTEMS

Lead : Sonia Kéfi and Jose Montoya

The growth of the human population and its standard of living leads to overexploitation of land and oceans, disrupting the global climate and leading species to extinction. The ecological responses to these disturbances are inevitably complex and require measures which allow them to be accurately described. Collectively, these measures assess the overall "stability" of ecological systems. Both disturbances to natural systems and the concept of stability are multidimensional. On the other hand, our understanding of these is not. This means that we have a remarkably poor understanding of the effects of different characteristics of the ongoing planetary changes on ecological stability.

The fragmented and one-dimensional approach adopted by environmentalists has led to ambiguities about the nature of stability as well as a clear disconnection between the theoretical and empirical literature on the subject (Donohue et al. 2016). Most of the theoretical work in ecology has been interested in the local asymptotic stability of ecological systems in the vicinity of an equilibrium and generally quantified this form of stability by the “resilience” of a system (Donohue et al. 2016, Kéfi et al. al. 2019), i.e. by the speed of its asymptotic return to equilibrium as given by the dominant value of the community matrix (May 1973). Most empirical studies, on the other hand, have used more practical measures of stability, such as the inverse of the coefficient of variation over time of the variables studied (Tilman et al. 2006, Hector et al. 2010).

A first question that arises is that of the relationship between the different stability measures used in ecology (Donohue et al. 2013): which metrics are redundant between them and which ones contain different information? In recent years, work has started to elucidate the relationships between certain stability metrics (Arnoldi et al. 2016, 2018, Arnoldi and Haegeman 2016, Dominguez-Garcia et al. 2019).

Another question is that of the relationships between biodiversity and different measures of stability, as well as the mechanisms responsible for these relationships. Recent empirical research has shown that biodiversity can indeed increase or decrease stability, depending on the measure of stability considered (e.g., resistance and variability) (Pennekamp et al. 2018).

The disturbances are also multidimensional. A recent study has shown that the type of disturbance can modify the relationship between biodiversity and stability. For example, even considering a single metric of stability, a change in biodiversity can increase or decrease the temporal invariability depending on whether the disturbance mainly affects the most abundant or least abundant species within a community (Arnoldi et al. 2019). Likewise, the type of habitat loss that affects food webs can increase or decrease their stability, as measured by the temporal variability of populations (McWilliams et al. 2019). If habitat is contiguously or correlated (i.e., large portions of habitat are lost while others remain intact), population variability increases. On the contrary, if habitat is lost at random in a given landscape, the variability of populations decreases.

In addition, most of the research on the relationships between biodiversity and ecosystem stability has been conducted primarily at the local level. However, the loss of biodiversity considerably reduces a number of ecosystem services by altering the functioning and stability of ecosystems at large temporal and spatial scales, which are most relevant for conservation and management (Gonzalez et al. In press, Isbell et al. 2015). A new theoretical study shows that biodiversity is becoming increasingly important for the functioning of ecosystems at larger spatial scales (Thompson et al. 2018). Likewise, the sign and intensity of the relationships between biodiversity and ecological stability change from one spatial scale to another. In particular, the temporal stability of a population or an ecosystem increases with the spatial scale (Wang and Loreau 2016, Wang et al. 2017, Delsol et al. 2018).

The objective of this GDR on this subject is threefold:
(1) establish the theoretical links between the different dimensions of stability in the face of different types of disturbances;
(2) develop theoretical models to study and better understand the relationships between the stability of ecosystems and their biodiversity;
(3) link this new theory to the experiments underway in France.

THEME 2. DYNAMICS OF INTERACTION NETWORKS

Lead : Elisa Thébault and François Massol

Interactions between ecological entities, whether they are interactions between individuals of the same species or different species, or interactions between populations of the same species structured in space (e.g. via dispersion), are central elements in ecology. Understanding these interactions requires integrating them into a network, that is, an object that includes all of the relationships (network links) that link a collection of entities (network nodes). Because they summarize important ecological information, interaction networks are the basis of many fundamental questions in ecology, whether it is to understand the mechanisms that determine interactions between species or to predict the consequences of these interactions on the dynamics. ecological systems and their responses to various disturbances (loss of species, etc.).

During the past five years of GDR TheoMoDive, the thematic group on interaction networks has been particularly active (discussions and presentations at each annual meeting, organization of a workshop on the theme of temporal networks). This group made it possible in particular to better identify emerging themes on interaction networks and to compare work on these themes with very diverse modeling methods, such as for example stochastic block models (SBM and LBM), joint species distributions (jSDM) or pattern frequency analyses in networks. The new version of the GDR's "Network of interactions" thematic group will continue in this vein by focusing on two themes that have experienced great growth in recent years in the study of networks.

1. Understand how multiple interactions in networks determine the dynamics of ecological communities and the functioning of ecosystems
So far, the vast majority of network studies have focused on given types of interactions, studied in isolation from each other: food webs (predation interactions), host-parasite networks, mutualistic plant-pollinator networks etc. Certain interactions and groups of species have also been ignored in empirical and theoretical approaches. Thus, most of the food webs studied ignore the interactions with the abiotic components of the ecosystem, while we know that many organisms (the so-called "ecosystem engineers" species) strongly modify the abiotic properties of their environment. Integrating this diversity of interactions into the description of ecological networks has been a major issue in the field of network research over the past five years (Kéfi et al. 2015, Pilosof et al. 2017, Astegiano et al. 2017, Hutchinson et al. 2019). This multiplicity of types of interactions in networks also questions classical theoretical approaches which generally consider only one type of interaction at a time (Bastolla et al. 2005). The few existing works on this subject show that taking into account various types of interactions (trophic, mutualist, involving the abiotic environment or the recycling of nutrients) modifies our understanding of the links between complexity and stability of communities (Mougi and Kondoh 2012, Sauve et al. 2014, Sanders et al. 2014) and affect the relationship between diversity and ecosystem functioning (Miele et al. 2019). In this context, the thematic working group will aim to develop theoretical research and models with a more integrative approach to the study of networks considering the diversity of interactions in ecosystems.


2. Understand the spatial and temporal dynamics of interaction networks
Empirical studies of networks of interactions have historically focused mainly on snapshot descriptions of networks at a given point, ignoring the spatial and temporal variability of interactions between species. The last few years have seen the development of empirical studies considering the spatial and temporal variability of interaction networks, and at different scales, e.g. intra-annual variability (CaraDonna et al. 2017), interannual (Ponisio et al. 2017), or between sites (de Manincor et al. in press, Kaiser-Bunbury et al. 2017). In this context, the objective of the working group will be to bring out new theoretical models allowing us to understand the temporal dynamics of networks, over short periods of time (from one year to the next, taking into account demographic fluctuations within communities and the effects linked to metacommunities) and over longer periods of time, in particular via the evolution of interacting species in a context of planetary changes. On this aspect, the group will seek to diversify its approaches by relying both on the development of deterministic (coupled differential equations in particular), stochastic (e.g. Bayesian network type) and qualitative models (use of formal grammars to describe the set of possibilities). The group will also seek to strengthen modeling approaches on spatially structured interaction networks (meta-networks and meta-ecosystems, e.g. Gravel et al. 2016), in particular to understand how the spatial structure of networks affects their functioning and their stability or resilience.

THEME 3. SPATIAL DYNAMICS, METACOMMUNITIES AND META-ECOSYSTEMS

Lead : François Massol and Isabelle Gounand

The concepts and models linking local dynamics and regional dynamics are defined by means of the prefix “meta-”, by analogy with the first of them, metapopulation (population of populations). A metapopulation (Levins 1970) is a spatial network where each node can host a population of an organism (Hanski and Gilpin 1997). By extension, a metacommunity designates a spatial network where each node can host a community (Leibold et al. 2004). When these species are structured, either as a food web, or more generally as an ecosystem, we will also speak of a meta-network or a metaecosystem (Loreau et al. 2003, Massol et al. 2011). The "meta" approach aims to answer classic questions such as the stability or coexistence of species within communities, but also more applied questions, such as the functioning of ecosystems, the emerging topology of interaction networks. and conservation biology. The concepts of metacommunity and meta-ecosystem have produced great advances in the understanding of the structuring of ecological systems at different spatial scales and are able to generate predictions of testable patterns in natura (Leibold et al. 2004, Logue et al. 2011).

Over the past five years, the "Metacommunities" working group within the GDR TheoMoDive has regularly brought together a highly active community, particularly around new approaches and interfaces with other disciplines (e.g. use of qualitative models for predict the possible states of communities on a dynamic landscape). The scope of the "meta" approach has greatly diversified, exploring the fields of the spatial ecology of food webs, ecosystem functioning and biogeography. A spatial approach to the functioning of ecosystems based on theories and concepts related to metacommunities and meta-ecosystems has thus found its place in the landscape of scientific ecology. For example, considering the dynamics of nutrient recycling no longer at the local scale, but at the scale of a meta-ecosystem, taking into account the flows of detritus and nutrients, leads to more theories and predictions. rich on the dynamics of these systems (Gravel et al. 2010a, 2010b, Gounand et al. 2014, Marleau et al. 2015), as well as the integration of food webs in a spatially structured system (Rooney et al. 2008, Gravel et al. 2011, Calcagno et al. 2011, Pillai et al. 2011). In addition, metacommunity approaches are beginning to integrate the genetic evolution of species (Urban et al. 2008). For example, the evolution of species can affect local competitive hierarchies and therefore niche effects according to the “monopolization” hypothesis (De Meester et al. 2002), or even modulate the effects of climate change on biodiversity via an evolution of the niche of competitively interacting species (Norberg et al. 2012).

The objective of this working group for the next five years will be to coordinate and stimulate theoretical research carried out in France in the field of metacommunities and meta-ecosystems in order to improve understanding of the spatial dynamics of biodiversity, food webs, ecosystem functioning, species evolution and the effects of global change. More specifically, we plan to support two particularly significant questions in the ecology of spatialized systems.

1. The effects of processes at work in meta-ecosystems on the structure and functioning of landscapes subjected to anthropogenic disturbances


In the context of global change, ecosystem transformations and the destruction of natural habitats have progressed considerably over the past three centuries. These changes in the environment are major factors in the current biodiversity crisis and the decline in ecosystem services (Fahrig 2003, Gonzalez et al. 2009). The spatially implicit framework proposed by "meta" approaches makes it possible to address the dynamics of such changes in fragmented habitats and thus to produce methodological tools adapted to conservation issues. For example, the ecology of metapopulations has allowed the emergence of concepts of minimum size and capacity for the viability of metapopulations (Lande 1987, Ovaskainen and Hanski 2001). The objective of the working group will thus be to bring out new theoretical work extending this approach to meta-ecosystems and taking into account a spatially explicit framework, e.g. via ecosystem networks or via continuous space approaches (partial differential equations); this extension of spatial ecology to the ecosystem level will make it possible in particular to explore how the dynamics of biological communities interact with the spatial heterogeneity of nutrients, how this interaction contributes to the maintenance of biodiversity, and how it modulates the response of biodiversity to anthropogenic disturbances.

2. The effects of planetary changes (in particular, climate change) on the distribution of interacting species, at ecological and evolutionary time scales.


The combination of ecological and evolutionary forces that determine the local coexistence of species and extrinsic limits to dispersal shape the present and future distribution of species. Classical approaches (SDM) predict changes in the distribution of species based on their niche and climate projections, but neglect the dynamics of interaction with other species (e.g: predation, facilitation, co-evolution) and with the environment (ex: niche construction, adaptation). The integration of these aspects in biogeography via the meta-ecosystem approach and the incorporation of evolutionary dynamics in meta-ecosystems will make it possible to refine these predictions by identifying the main mechanisms of the extinction-colonization dynamics at the borders of the distributions of species.

THEME 4. DYNAMICS OF INTERACTIONS BETWEEN BIODIVERSITY AND SOCIETY

Lead : Michel Loreau, Cédric Gaucherel and Sébastien Barot

Historically, ecology has primarily started by studying the dynamics and functioning of populations, communities and natural ecosystems. As a result, the theoretical corpus of ecology is essentially based on ecological processes relatively independent of human activities and societies. However, it has become increasingly clear that these have a significant and globally negative impact on biodiversity (Pereira et al. 2010) and on the functioning of ecosystems and the biosphere (Vitousek et al. 1997, Cardinale 2012), in particular because of changes in land use, global warming and the exploitation of certain species. This observation has led, little by little, to the study of new ecosystems created or modified by humans (Hobbs et al. 2009), such as urban ecosystems (Barot et al. 2019) or agroecosystems (Wezel et al. 2009), just like natural ecosystems. At the same time, a current of research has developed around the concept of ecosystem services, their evaluation and their use in decision support (Costanza et al. 1997, Gomez-Baggethun et al. 2010). This research program has helped to show that human societies depend on biodiversity and the functioning of ecosystems and that it is therefore necessary to adapt our interactions with biodiversity so as not to threaten the services it provides us and to preserve the sustainability of our companies. In doing so, biodiversity and other components of ecosystems will often benefit from less impact. In other words, it is no longer just a question of studying ecological systems; it is also necessary to analyse socio-ecosystems and all the feedback mechanisms between biodiversity, physical and chemical processes and human societies. It is in part on this conceptual framework that the work of IPBES is based (Diaz et al. 2019).

Such an objective requires not only new empirical approaches describing the feedback mechanisms between human societies and biodiversity, but also and above all new developments in theory and modeling. On the one hand, predicting the dynamics resulting from these complex feedbacks cannot be done without new mathematical or computer tools. On the other hand, these tools will have to take into account mechanisms absent from models conventionally developed in ecology: for example, psychological, social, economic and political mechanisms leading to decisions and actions that influence biodiversity and ecosystems. For these reasons, several philosophers and ecologists propose to see the ecosystem as a whole, without particular polarization between humans and non-human components (Naess 1975, Gaucherel and Pommereau 2019). It requires considerable conceptual and technical development to answer many original scientific questions. For example, we can ask ourselves whether the taking into account of human societies leads to qualitatively original dynamics compared to the ecological dynamics traditionally studied, or again, what are the conditions for these feedbacks between societies and biodiversity to lead to a balance.

Opening up this field of research involves new theoretical research, often interdisciplinary, in which the GDR wishes to participate. This research includes modeling work focusing on a particular feedback mechanism (Martinet et al. 2007), possibly on a local scale, and more integrative models and on a global scale (Henderson and Loreau 2019), even qualitative approaches. inspired by theoretical computer science (Gaucherel and Pommereau 2019). They aim to analyse, understand and predict the dynamics of socio-ecosystems, but also have an important role to play in resolving the current environmental crisis, by shedding light on the range of possibilities and proposing possible solutions in line with the science of sustainability (Clark and Dickson 2003).

THEME 5. THE ROLE OF EVOLUTION

Lead : Emanuel Fronhofer and Florence Debarre

Ecology and evolution have long remained separate disciplines. The study of the dynamics of communities or food webs is often complex enough not to add an evolutionary component; conversely, in evolutionary biology, the dynamics of populations and communities are often ignored or simplified, in particular, in the case of theoretical models, to allow a mathematical analysis.
Recently, however, connections between ecology and evolution have multiplied. In France, in particular, this thematic merger has resulted in a visible rapprochement of communities of researchers in ecology and evolution through the change of the French Society of Ecology (sfe) into the French Society of Ecology and Evolution (sfe2) and a series of workshops, such as the Symposium `` At the border between ecology and evolution '', organized by Sfe2 and LabEx CeMEB in Montpellier.

The role of evolving ecology and the role of evolution in ecology are therefore increasingly taken into account, in line with historical models in evolutionary ecology. These models were particularly interested in feedbacks between demography and evolution (single-species models), the effect of spatial structure, but also interactions between species (competition, predation, parasitism, mutualism) and between communities (Govaert et al. 2019). Models explicitly representing interactions with the environment can also include eco-evolutionary dynamics (e.g. Estrela et al. 2019).

These models, however, are most often fairly simple and include little biodiversity (often only two or three species are modelled by a system of ordinary differential equations), although there are exceptions (Govaert et al. 2019). It is therefore high time to ask in a more concrete way the question of how evolution affects the dynamics of biodiversity. This question is particularly relevant since most of the global changes that directly threaten biodiversity and the functioning of ecosystems are also important selective forces.
It is therefore necessary to develop powerful approaches that allow us to understand the functioning of ecosystems in an evolutionary context. In their review article, Govaert et al. (2019) recommend in particular integrating more realism into eco-evolutionary models (for example, to consider the effect of plasticity or the structure of populations), but also to adopt mechanistic approaches, in particular by explicitly considering interactions between individuals (instead of phenomenological approaches). Another methodological challenge is to consider eco-evolutionary dynamics occurring on comparable time scales, instead of the temporal decoupling (useful for mathematical analysis) traditionally assumed.

The aim of the GDR is to coordinate theoretical developments, to create a network of researchers and to organize workshops on this topic. This theme can also be considered as cross-cutting within the GDR, with development issues affecting all subjects.

Transversal approach

TRANSVERSAL APPROACH 1. DATA-THEORY LINK

TRANSVERSAL APPROACH 2. BEYOND EQUILIBRIUM

TRANSVERSAL APPROACH 3. GENERAL LAWS IN ECOLOGY

TRANSVERSAL APPROACH 1. DATA-THEORY LINK

Lead : Frédéric Barraquand and Arnaud Sentis

The coupling of theoretical models and empirical data makes it possible to strengthen our understanding of ecological processes by contrasting alternative ecological hypotheses formalized mathematically and by proposing a theoretical framework for the interpretation of ecological data. Nevertheless, although the first ecological theories relied heavily on empirical data (Kingsland 1995), a divide between theory and data has developed over the years, particularly during the second half of the last century. This has led to a specialization and a gap between theoretical ecologists, living in a world of Lotka-Volterra models and others, and specialists in field or experimental studies. Overcoming this cleavage would enrich models and theories by making them more realistic and strengthen empirical approaches by providing them with a theoretical framework with hypotheses to be tested. Faced with this observation, efforts have been made and the cleavage has been greatly reduced in recent years (Kendall 2015), which offers substantial opportunities for theorists to make more predictive theoretical models (Dietze et al. 2018) and thus increase our ability to falsify theoretical predictions. Moreover, beyond the production of hypotheses explaining the phenomena, the theory plays an important role in the clarification of the concepts and the meaning of the measured quantities, i.e., the theory makes it possible to verify that what one measures is what we want to measure (e.g Berlow et al. 2004). As part of the GDR, it is therefore natural to focus on approaches that allow better coupling between theory and data.


A promising approach is the development of methods for estimating the parameters of mechanistic models from experimental data such as time series (Ionides et al. 2006, Rosenbaum et al. 2019, Pennekamp et al. 2019). These approaches make it possible to calibrate theoretical models in order to then test their predictive power or to contrast various hypotheses on the functioning of a system, for example to explain fluctuations in the abundance of species (e.g. Kendall et al. 2005). Inference methods are used to fit systems of differential equations to time series derived from laboratory experiments or empirical data (Ellner et al. 2002, Rosenbaum and Rall 2018). The use of these methods is facilitated by the increase in the digital computing power of computers and opens up real prospects for theory-data coupling. Nevertheless, many questions remain, in particular on:
- the ability of the methods to recover the parameters of the theoretical models (i.e., identifiability);
- the type of data to be collected in the field;
- the architecture of the models to be adjusted to the data.
A particularly relevant project is the development of methods which make it possible to combine the non-linearity of ecological processes with stochasticity, which is omnipresent in ecology (e.g., Ionides et al. 2006).

As part of the study of the coexistence of competing plant species, a use of theory well connected to the data has shown both which quantities to measure in the field (Hart et al. 2018), which mechanistic models to adjust data to contrast ecological hypotheses (Adler et al. 2010), and how to reconcile the apparently contradictory results of experiments with those of in situ measurements (Tuck et al. 2018).

These few examples show that the links between theory and data apply to a set of thematic fields, hence the transversal aim of this axis in the GDR. The thematic working group devoted to this theme intends to focus its reflection on the issues of connection to empirical data, beyond the simple data-model adjustment, in particular through discussions and review articles. By bringing together ecologists of different backgrounds around the use of data in theoretical ecology, this activity of the GDR will make it possible to seek the consensus on which ecologists can agree and to explore the approaches which make it possible to strengthen the links between theory and data.

TRANSVERSAL APPROACH 2. BEYOND EQUILIBRIUM

Lead : Claire de Mazancourt

One of the recurring criticisms of the theory is about the commonly used assumption that systems are at or near equilibrium. However, the real systems observed seem to be much more dynamic (Lundberg et al. 2000, Inchausti and Halley 2003). The equilibrium hypothesis can be useful because it allows for the in-depth study of a system from a mathematical point of view, but it can also seem quite simply incompatible with field observations. The notion of stationary state, with distributions of states distributed with certain frequencies, was developed to overcome this problem, but disturbances and natural systems are not always stationary. To study systems out of equilibrium, theoretical tools exist and are being developed within the GDR Theory and Modeling of Biodiversity and the international scientific community of theoretical ecology. There are several factors that can push a system out of balance. Each has a different theoretical treatment.

1. Transient dynamics
The experiments carried out simply do not have time to reach equilibrium, hence the interest in the theory of studying transient dynamics on the way to equilibrium. Thus, the short term response of a system is very different from the long term response. As most of the theory concerns the long-term response, the development of a theory of short-term responses is particularly welcome (Arnoldi et al. 2018).

2. Fluctuations of extrinsic origin
More profoundly, systems might not tend to a steady state. When it comes to extrinsic fluctuations, one expects the existence of a state of equilibrium but which changes over time as the external conditions change, stochastically, cyclically (seasons) or gradually (change directional, such as climate change). Concepts and techniques developed by mathematicians in the theory of non-autonomous dynamics have recently been introduced in ecology (Chesson 2017) to study these phenomena.

3. Fluctuations of intrinsic origin
Fluctuations such as cycles and chaos can emerge in very simple systems. They have typically been studied in low dimensional systems (May 1976). Maintaining chaos in complex systems requires specific conditions and techniques to detect them (Roy et al. 2019).

This transversal approach raises the question of the link between theory and empirical data: what is the prevalence of ecological systems that are out of equilibrium? Is the theory still applicable, even on non-equilibrium systems? Should the theory be adapted to non-equilibrium systems? Can the intrinsic and extrinsic factors which cause this out of equilibrium dynamic be demonstrated and quantified?

To conclude, this transversal approach will aim to research and develop existing tools to study non-equilibrium systems and related issues. It will also push researchers at GDR to question the relevance of using these tools or developing new ones.

TRANSVERSAL APPROACH 3. GENERAL LAWS IN ECOLOGY

Lead : Cédric Gaucherel and Sébastien Barot

It is not common to question the laws of ecology; Yet this is exactly what this working theme is proposing within the GDR. What do we mean by that? A detour through neighbouring disciplines can enlighten us. With at least four centuries of hindsight, physics has a long tradition of perfecting laws, making it possible to explain in a reliable and universal way (by definition, proven and without exception) its phenomena (Barberousse et al. 2000, Hartmann and Frigg 2005). This is not the case with ecology and evolutionary sciences, and even with proven principles (by definition, which are not questioned) of theories (sets of principles) such as natural selection or Mendelian laws are still much debated (Mayr 2004, Morange 2017). Of course, models regularly challenge accepted principles and test their associated hypotheses (Israel 1996, Hartmann and Frigg 2005).

Biology is readily quite reductionist, ecology, more holistic, sometimes seeks universal laws (Hubbell 2001, Brown et al. 2002, Gaucherel 2013) and continually questions the reliability of the principles that it brings to light, even on the universality of the theories which could result from it. Whether or not it is believed that there are laws in ecology or not, several interesting corollary questions emerge from this question. For example, one is entitled to ask whether a law should relate to mechanisms (processes) or reasons (patterns)? Because while physical laws are generally based on mechanisms, some of them (e.g. gravitation) often resemble patterns and retain a phenomenological flavour (e.g. fractals and allometries). Some even question what the mechanisms are and what any causal explanation is worth (Israel 1996, Barberousse et al. 2000). And when ecology highlights allometric laws, it is part of this same tradition (Brown et al. 2002, Hatton et al. 2015). In addition, some laws are purely statistical (e.g. law of large numbers), far from any mechanism (Frank 2009).

And if ecologists highlighted a law in their field, would it have ecological specificities? In particular, biology and ecology all know the importance of scales and levels of organization in living things. Would an ecological law be affiliated with a particular level of organisation? Certain laws well known to ecologists attempt to explain the distribution of species, either by chance (Hubbell 2001) or by other observed regularities (Neill and Gignoux 2008, Tilman 2011). Several of them link several ecological compartments or several levels of organisation between them (Hardin 1960, MacArthur and Wilson 1963, Tilman 2011). Can these principles be considered as laws? We can guess that the status of such laws will not be similar to that of physical laws which appear, in the eyes of biologists, much more robust and universal (Putnam 1975, Gaucherel 2013). What exceptions, what variations, in these ecological laws, are we ready to accept? And further, how to manage the contradictions or the possible interactions between ecological laws?



Ecology does not have to follow the path of physics or chemistry, but it is entitled to be inspired by them. It can also be inspired by all related disciplines, from physics (Brown et al. 2002, Neill and Gignoux 2008), to economics, including linguistics (Gaucherel 2019). Several recent attempts go in this direction, and a field of research illustrates this growing interest in laws in ecology. Indeed, it would be daring to claim that ecological objects and phenomena violate physical laws, such as thermodynamics, or biological laws, such as natural selection (Harte 2002). Beyond the laws themselves, it is the history of neighbouring disciplines, particularly biology, that can prove to be a source of innovation in ecology (Morange 2017). The old debates on evolution, a field which made a noticeable irruption in ecology, show that it is interesting to seek laws where, previously, one still had only a list of processes and a classification. objects.
Still, the more pragmatic would question: What would be the use of such laws? Would they be there for the sole intellectual pleasure of theorists? Or would they have a real daily utility for managers, farmers or geneticists? Would such laws help to make predictions? Many think so, and claim that a better knowledge of the mechanisms and principles associated with central objects, such as the ecosystem or the population (Hardin 1960, Tilman 2011), would undoubtedly provide assistance to those who have to manage these objects on a daily basis. still poorly understood (Gaucherel 2019). Even if it turned out that ecology does not have laws, reflecting on this question would contribute as much to basic ecology as it does to applied ecology.
As part of this transversal axis we will interact with philosophers of science and we want to design a first list of general laws by doing an online survey.


Complementarity with the GDR Statistical Ecology


The GDR Statistical Ecology aims to develop and disseminate statistical methodologies adapted to the various fields of ecology, including in particular the algorithmic aspects of the fit to data (maximum likelihood, Markov Chain Monte Carlo), data management (data citizen, ...) and conceptual of applied statistics (e.g, frequentist vs Bayesian philosophy). The GDR Theory and Modeling of Biodiversity addresses questions upstream and downstream of the use of these methodologies: which data to seize for which questions? Which statistical models also make theoretical sense / how to theoretically interpret their parameters? Are there classical empirical analyses that the theory can prove are uninterpretable? How to reduce the complexity of theoretical models to fit them to necessarily limited data? Are there ecological questions of importance to the theory that currently escape empirical analysis?

Participating laboratories

  • Station d’Ecologie Théorique et Expérimentale

    Moulis (UMR 5321)

    • Ecologie du Paysage et Conservation de la Biodiversité
    • Théorie et Modélisation de la Biodiversité

  • Laboratoire Evolution et Diversité Biologique

    Toulouse (UMR 5174)

    • Diversification Évolutive et Écologique des Populations
    • Ecologie Aquatique et Changements Globaux

  • Institut d’Ecologie et des Sciences de l’Environnement

    Paris (UMR 7618)

    • Ecologie et Evolution des Réseaux d'Interactions
    • Ecologie Intégrative: Des Mécanismes aux Services Écosystémiques

  • Institut de Biologie de l’Ecole Normale Supérieure

    Paris (UMR 8197)

    • Eco-Evolution Mathématique
    • Modélisation de la Biodiversité

  • Laboratoire d’Ecologie Alpine

    Grenoble (UMR 5553)

    • Bases Génétiques de l’Adaptation
    • Evolution, Modélisation et Analyses de la Biodiversité
    • Fonctionnement, Dynamiques et Services des Ecosystèmes de Montagne

  • Institut des Sciences de l’Evolution

    Montpellier (UMR 5554)

    • Dynamique de la Biodiversité
    • Ecologie et Evolution des Communautés

  • Laboratoire Biodiversité Marine et ses Usages

    Montpellier (UMR 9190)

    • Dynamique et Fonctionnement des Communautés

  • Unité de Recherche Ecologie Fonctionnelle et Biogéochimie des Sols et des Agroécosystèmes

    Montpellier (UMR 210)

    • Nutriments et Intensification Écologique
    • Sols, Activités et Réseaux Biologiques

  • Centre d’Ecologie et des Sciences de la Conservation

    Paris (UMR 7204)

  • Laboratoire Evolution, Ecologie, Paléontologie

    Lille (UMR 8198)

    • Evolution et Ecologie

  • Centre de Mathématiques Appliquées

    Palaiseau (UMR 7641)

  • Laboratoire d’Ingénierie pour les Systèmes Complexes

    Clermont-Ferrand (IRSTEA)

  • Unité de Recherche Écosystèmes Montagnards, Grenoble

    Grenoble (IRSTEA)

    • Dynamiques et Gestions des Écosystèmes de Montagne

  • Centre d’Ecologie Fonctionnelle et Évolutive

    Montpellier (UMR 5175)

    • Dream
    • Dynamique des Paysages et de la Biodiversité
    • Ecopar
    • Génétique et Ecologie Évolutive

  • Institut Sophia Agrobiotech

    Sophia Antipolis (UMR 1355)

    • Ecologie Théorique et Appliquée aux Agrosystèmes semi-confinés

  • LabEx COTE

    Université de Bordeaux

    • Theoretical and Integrative Ecology

Scientific Council

The Scientific Council of GDR TheoMoDive is composed of:

  • Sébastien Barot - Institut d’Écologie et des Sciences de l’Environnement, Paris (UMR 7618)

  • Jérôme Chave - Laboratoire Évolution et Diversité Biologique, Toulouse (UMR 5174)

  • Claire de Mazancourt - Station d’Écologie Théorique et Expérimentale, Moulis (UMR 5321)

  • Franck Jabot - Laboratoire d’Ingénierie pour les Systèmes Complexes, Clermont-Ferrand (IRSTEA)
  • Sonia Kéfi - Institut des Sciences de l’Évolution, Montpellier (UMR 5554)
  • Michel Loreau - Station d’Écologie Théorique et Expérimentale, Moulis (UMR 5321)
  • François Massol - Laboratoire Evolution, Ecologie, Paléontologie, Lille (UMR 8198)
  • José Montoya - Station d’Écologie Théorique et Expérimentale, Moulis (UMR 5321)
  • Hélène Morlon - Institut de Biologie de l’École Normale Supérieure, Paris (UMR 8197)
  • Nicolas Mouquet - Institut des Sciences de l’Évolution, Montpellier (UMR 5554)
  • Elisa Thébault - Institut d’Écologie et des Sciences de l’Environnement, Paris (UMR 7618)
  • Wilfried Thuiller - Laboratoire d’Écologie Alpine, Grenoble (UMR 5553)

Grants

The GDR TheoMoDive offers, within the means available, grants for doctoral students and postdoctoral researchers who wish to do a research stay in one of the participating laboratories.


Applications for grants should be addressed to Claire de Mazancourt -  This email address is being protected from spambots. You need JavaScript enabled to view it. and contain:

  • a CV

  • a brief description of the objectives of the proposed stay

  • a brief justification of the planned expenditure.

These requests are then examined by the Scientific Council of the GDR.

Annual meetings, Workshops and Seminars

Annual meetings


The annual meetings make it possible to ensure the continuity of the reflections and work carried out within the GDR, to synthesize them and to identify new avenues of research. Their objective is less to draw up an exhaustive assessment of the work carried out than to stimulate common reflection.
As a result, they focus on new advances made, new developments underway and new projects in the pipeline, as well as on cross-cutting issues across GDR.

Workshops


The GDR supports, within the means available, smaller workshops to advance reflections and research on the various research themes of the GDR.
Workshops at the interface of several themes or allowing new themes to emerge are also encouraged.

Workshop proposals should be sent to Claire de Mazancourt - This email address is being protected from spambots. You need JavaScript enabled to view it. and contain:

  • a description of the objectives of the workshop

  • a list of potential participants

  • a provisional program

These proposals are then examined by the Scientific Council of GDR.

Seminars :

The new webinar series of the GdR TheoMoDive will start on Tuesday March 29 at 3pm CET with a talk by Jean-François Arnoldi (SETE-Moulis) on "Invasion- and extinction-induced abrupt shifts in ecological communities: the role of fitness and feedbacks" who represents the GdR's reseaech theme "Diversity, stability, and functioning of ecological systems". The Zoom link will be send around via the GdR mailing list.

Mailing List

Information regarding GDR activities, in particular annual meetings and workshops, is disseminated via the mailing list: This email address is being protected from spambots. You need JavaScript enabled to view it.

  • To subscribe: Send a message This email address is being protected from spambots. You need JavaScript enabled to view it. from the address from which you wish to subscribe with the subject: subscribe gdr-theomodive Your First Name Your Last Name

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Contact

GDR TheoMoDive
CNRS - SETE (Station d'Ecologie Théorique et Expérimentale)


2 route du CNRS
09200 MOULIS


Contact : Claire de Mazancourt - This email address is being protected from spambots. You need JavaScript enabled to view it.

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Thèmes de recherches transversales

Thèmes de recherches

Transversales

Lier les processus à plusieurs échelles : du local au global (metascales)

Il est reconnu que la structure spatiale des paysages joue un rôle crucial dans la dynamique écologique et évolutive. Les organismes interagissent entre eux et avec leur environnement local, avec des schémas à l'échelle régionale. Notre compréhension des mécanismes à l'origine de ces interactions à l'échelle locale est essentielle à notre compréhension de la dynamique des populations, des communautés et des écosystèmes. Cependant, si ces connaissances peuvent être étendues à des modèles régionaux et mondiaux, ou si d'autres mécanismes sont importants à ces échelles supérieures, il faut de nouvelles recherches.

L'objectif de cet axe transversal est de rassembler des chercheurs travaillant à différentes échelles (des individus aux écosystèmes et à la macro écologie) pour identifier les éléments méconnus dans notre compréhension de cette problématique, et contribuer à combler ces connaissances par le développement d'un projet commun.

Pour aborder ce sujet, nous réunissons plusieurs approches et des expertises pour intégrer la théorie, les expériences et les analyses de ‘big data’ à plusieurs échelles.

Les thèmes comprennent :

(1) le passage des populations aux méta-écosystèmes,

(2) l'intensification de la recherche sur la biodiversité locale, la structure, la fonction et la stabilité du réseau à des étendues spatiales plus vastes, et

(3) l'étude des modèles mondiaux de la biodiversité et du fonctionnement des écosystèmes.

Responsables :

Jose Montoya et Staffan Jacob

Expérimentation, observation et causalité

L'utilisation d'expériences en écologie et évolution a augmenté de façon spectaculaire au cours des deux ou trois dernières décennies, comme en témoigne la construction récente de nombreuses installations expérimentales hautement contrôlées et la multiplication d'expériences à long terme dans les écosystèmes naturels. Bien que certaines revues aient déjà discuté de l'utilisation des installations expérimentales et des connaissances scientifiques qui peuvent en être tirées, il n'y a eu aucune tentative complète de revoir le rôle des expériences en écologie, leurs contraintes et limites, et la façon de surmonter ces dernières (temps, espace, environnements extrêmes, facteurs de stress multiples, nouvelles perspectives d'expérimentation, combinaison d'expériences, d'observations et de modèles, etc.).

La SETE est bien placée pour lancer une réflexion sur ces questions. Ainsi nous abordons les problématiques suivantes :

  • l'utilisation de systèmes modèles dans les expériences écologiques et leurs limites ;
  • reproductibilité et validité des expériences écologiques ;
  • statistiques et causalité ;
  • rétroactions entre expériences écologiques, théorie écologique et modélisation écologique;
  • rétroactions entre expériences et observations dans les systèmes naturels ;
  • expériences et échelle : combien de temps est assez ? Quelle taille est assez grande ?
  • dynamique évolutive des écosystèmes complexes ;
  • expérimentation des rétroactions entre l'écosystème et l’homme ;
  • expériences sur les mosaïques des écosystèmes ;
  • limites des expériences /facteurs multiples et étude des événements extrêmes.

Responsable :

Jean Clobert

Retours éco-évolutifs

Les rétroactions entre les dynamiques écologiques et évolutives sont de plus en plus reconnues comme importantes. Elles sont étudiées à la SETE en utilisant différents systèmes de modèles (protistes, microbiote, papillons, poissons, écrevisses, libellules) et une combinaison d'approches (observationnelles, expérimentales, modélisation). L’une des priorités est la dispersion, un trait-clé de la réponse d’un organisme aux changements globaux, avec de forts effets sur la dynamique écologique et évolutive. La dispersion évolue rapidement, montre une variation génétique au sein des métapopulations et module les relations entre les paramètres évolutifs et écologiques. La fragmentation naturelle et anthropique de l'habitat affecte l'évolution de la dispersion, dont l'influence sur la dynamique des populations peut se répercuter vers le haut sur la dynamique des communautés et le fonctionnement des écosystèmes. A la SETE, la dispersion est étudiée en identifiant et en mesurant plusieurs traits potentiellement corrélés qui forment des syndromes de dispersion héréditaires. Ce processus permet à la modélisation de faire des prédictions qui peuvent être testées par la libération expérimentale d'animaux dans des microcosmes et des métatrons terrestres et aquatiques.

Un deuxième axe de recherche sur les dynamiques éco-évolutives à la SETE concerne la diversité intraspécifique.

Il a été démontré que les changements dans la diversité intraspécifique génèrent des rétroactions évolutives agissant sur la dynamique écologique, y compris les effets au niveau de l'écosystème. Par exemple, l'évolution de la diversité fonctionnelle intraspécifique au sein des populations d'écrevisses influence le taux de décomposition des litières dans leur habitat lacustre.

Les recherches à la SETE montrent également comment les dynamiques écologiques et évolutives se combinent pour influencer des processus aussi divers que l'assemblage du microbiome des éponges, les probabilités d'extinction des populations de papillons et l'imbrication, la diversité et la résilience des réseaux mutualistes.

Responsables :

Claire de Mazancourt and Michael Singer

Interactions homme-nature

L'augmentation globale de la taille de la population humaine, la conversion des terres indigènes en agriculture et l'étalement urbain ont tous conduit à une fréquence et une intensité plus élevées des interactions entre l'homme et la nature. Les recherches menées à la SETE nous éclairent sur les problèmes qui affectent différents secteurs de la société. Il s'agit notamment de la planification de la conservation, de l'amélioration des services écosystémiques par une meilleure compréhension des rétroactions entre les activités humaines et le fonctionnement de l'écosystème, élucider la psychologie sous-jacente du comportement social et informer les politiques au niveau international sur la réponse à l'évolution de la biodiversité et du changement climatique. Les conséquences inattendues d'une augmentation des interactions homme-nature, telles que le transfert du virus VIH de l'animal aux populations humaines, peuvent avoir des effets dévastateurs. Bien que les effets négatifs de la production alimentaire intensive génèrent des pressions sociales pour la durabilité, les recherches à la SETE ont montré comment les retards de réponse à ces pressions peuvent, par leur influence sur les services écosystémiques, augmenter la vulnérabilité des systèmes socio-écologiques et entraîner plus probablement l'effondrement des populations humaines. Les recherches à la SETE ont mis en évidence l'importance de coordonner les études sur la dépendance humaine à la biodiversité avec les études des influences humaines sur la biodiversité.

La SETE est bien placée pour approfondir ces thèmes. Une question primordiale que nous développons est : quelles sont les implications de la pression croissante des moteurs du changement global sur les interactions entre les populations humaines et les systèmes naturels, et pour les processus de rétroaction? Des exemples de types de projets sur le thème général des interactions homme-nature :

(1) À l'échelle mondiale, le changement climatique anthropique et le transport humain de plantes et d'animaux exotiques entraînent des mouvements d'espèces sauvages et humaines à travers le monde. Ces changements dans les emplacements géographiques engendrent de nouvelles interactions au sein des communautés naturelles et humaines, et entre les humains et les espèces sauvages. Les questions émergentes que les chercheurs de SETE commencent à aborder sont : quels sont les impacts actuels et futurs de ces nouvelles interactions sur la biodiversité locale, la santé humaine, l'accès et la production alimentaire et le bien-être humain ? Comment une meilleure compréhension de la base génétique vs plastique du comportement coopératif, et de son évolution culturelle et génétique, peut-elle éclairer les actions politiques visant à modifier le comportement du public, par ex. réduire les émissions de gaz à effet de serre ou protéger la biodiversité ?

(2) À l'échelle régionale, les humains empiètent de plus en plus sur les zones naturelles environnantes, créant des habitats fragmentés et dégradés qui résistent moins aux perturbations naturelles ou à l'augmentation de l'intensité des perturbations humaines. À leur tour, les paysages fragmentés et dégradés sont moins adaptés pour accueillir à la fois les espèces sauvages et les humains qui sont chassés de leurs terres natales. Comment une meilleure compréhension des processus se produisant lors de la formation de nouvelles communautés sauvages et humaines peut-elle aider à améliorer la résilience et la stabilité des systèmes humains-naturels couplés ?

Responsable :

Camille Parmesan

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CHANGE- Axes de Recherche

CHANGE

Axes de Recherche

Impacts des génomes et des phénomes sur le fitness

Les traits phénotypiques et les gènes sont au cœur des études éco-évolutives. Cependant, il est bien connu que la sélection peut opérer simultanément sur les gènes et les traits dans des directions différentes et avec des forces variables au sein d'une population, ce qui limite notre capacité à interpréter correctement les modèles évolutifs.

Notre équipe développe une vision inclusive des phénomes et des génomes pour mieux comprendre les schémas adaptatifs. D'une part, nous décrivons la structure des traits phénotypiques dans divers taxons et leur potentielle organisation dans des syndromes, c'est-à-dire des suites de traits corrélés. Nous relions ces stratégies phénotypiques aux avantages ou aux coûts potentiels sur le fitness des organismes. La comparaison d'espèces étroitement apparentées nous aide à aborder ces questions aux niveaux intra- et interspécifique.

Notre équipe étudie notamment les syndromes de pollinisation chez les orchidées, les syndromes de prédation chez les serpents et les syndromes de dispersion chez les lézards et les protistes. Nous étudions également une série de traits complexes à travers toutes nos espèces modèles comme la socialité, la coopération, la communication, le mouvement, la capacité cognitive, la sénescence, la préférence d'habitat et l'organisation cellulaire. D'un autre côté, nous étudions l'impact de la structure du génome sur les corrélations entre caractères phénotypiques et sur le fitness. Deux systèmes modèles sont particulièrement utilisés étudiés : les éléments transposables chez les orchidées et la structure singulière du macronoyau des Ciliés (par exemple, nombre élevé et variable des copies chromosomiques).

Notre objectif à long terme est d'établir des cartes multidimensionnelles des génomes et des phénomes, mettant en lumière les corrélations entre un certain nombre de leurs dimensions.

Interactions entre les différents processus éco-évolutifs dans des environnements changeants

La réponse de la biodiversité aux changements environnementaux est une question scientifique et sociétale majeure en raison du changement global. Cependant, nous sommes loin d’être capables de prédire avec précision l’impact des changements environnementaux sur la persistance des organismes, car il nous manque une vision globale des interactions entre les divers processus à l’œuvre.

L’équipe évalue comment la sélection sur des variations préexistantes ou nouvelles, la plasticité phénotypique et/ou la dispersion permettent aux organismes de s’adapter aux changements environnementaux à court (d’une à quelques générations) et à long terme (des dizaines à des milliers de générations). Nous mesurons l'aptitude des organismes dans un ensemble de conditions naturelles ou expérimentales bien contrastées. En particulier, nous nous concentrons sur les réponses aux gradients environnementaux dans des études de terrain à long terme dans les Pyrénées et les Cévennes pour analyser, par exemple, la variation naturelle de la réponse des organismes à l'altitude, à l'hypoxie, à la température et à l'utilisation des terres.

Nous étudions également la fragmentation de l'habitat, le changement climatique, la pollution et leurs interactions en réalisant des expériences dans la nature et dans les installations expérimentales de la SETE (métatron terrestre, microcosmes).

Bases moléculaires de l’adaptation

La capacité des organismes à exprimer des phénotypes correspondant à des environnements changeants est directement liée aux mécanismes moléculaires sous-jacents.

Les modifications génétiques, mais aussi non génétiques, peuvent entraîner des changements phénotypiques qui permettent aux organismes de suivre les changements environnementaux. Lier l'évolution des phénotypes complexes à leurs bases moléculaires fournit ainsi des informations clés sur le temps de réponse et la stabilité de l'adaptation des organismes. Nous utilisons des approches fondées sur les génomes complets, les transcriptomes et/ou les épigénomes pour relier les modifications phénotypiques, fixes ou plastiques, à leurs mécanismes moléculaires sous-jacents.

Des modifications épigénétiques ou des changements de ploïdie (dans le macronoyau des ciliés) pourraient notamment permettre aux génotypes sous-optimaux d'atteindre rapidement un optimum de fitness par des changements plastiques, étape possiblement suivie par de l'assimilation génétique. Nous nous concentrons sur les déterminants moléculaires des traits influençant fortement le fitness, par exemple, la cognition chez les oiseaux ou l'organisation ciliaire chez les ciliés.

Nous utilisons l'évolution expérimentale pour étudier la dynamique des modifications génétiques et non génétiques. Nous recherchons notamment l'existence de contraintes (épi)génétiques responsables de l'évolution des syndromes phénotypiques chez les serpents, les orchidées et les ciliés.

Rôle de la variabilité intraspécifique sur la dynamique et le fonctionnement des populations, des communautés et des écosystèmes

Le fonctionnement et la dynamique des systèmes biologiques peuvent être très différents selon les assemblages de phénomes et de génomes. De plus, certaines stratégies telles que la croissance rapide, les interactions strictes entre espèces et de fréquentes insertions génomiques peuvent assurer un succès à court terme, mais une vulnérabilité à long terme en fonction de la dynamique globale des populations, des communautés ou des écosystèmes.

Nous étudions l'impact de la variabilité intra- et inter-spécifique sur la dynamique des systèmes biologiques à travers des échelles de temps écologiques et évolutives. Nous relions la démographie, la productivité de l'écosystème et la diversification des espèces à la variabilité du phénome et/ou du génome. Nous développons notamment des expériences concertées sur de multiples taxons entre laboratoires et pays pour que les réponses obtenues gagnent en généralité.

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Transversal Research Themes

Transversal

Research Themes

Linking processes across multiple scales: from local to global (Metascales)

The spatial structure of landscapes is widely recognized to play a crucial role in ecological and evolutionary dynamics. Organisms interact among them and with their environment at the local scale, with movements among localities linking patterns at the regional scale. Our understanding of the mechanisms driving these interactions at the local scale is key to our understanding of population, community and ecosystem dynamics. However, whether this knowledge can be scaled up to regional and global patterns, or whether other mechanisms play at these higher scales requires novel research. The objective of this transversal axis is to gather researchers working on multiple scales (from individuals to ecosystems and macroecology) to identify key gaps in our understanding of this issue, and contribute to filling these gaps by the development of a common project.

Addressing this topic will require bringing together the diversity of approaches and expertise present in SETE to integrate theory, experiments and big data analyses at multiple scales. Themes will include: (1) moving from populations to meta-ecosystems, (2) scaling-up research on local biodiversity, network structure, function and stability to larger spatial extents, and (3) studying global patterns of biodiversity and ecosystem functioning.

This transversal axis will be organised around activities at three timescales:

(1) Bimonthly internal meetings at SETE to strengthen interactions across teams through paper discussions and thematic topics. External seminars will be organised to further diversify the conceptual inputs of the discussions. From these regular meetings, we expect to reach strengthened day-to-day research interactions across teams, and to identify gaps and key issues we should focus on to provide breakthroughs.

(2) Annual workshops (2-3 days duration) on well-identified topics emerging from the regular meetings of the axis. These workshops will gather internal researchers and external speakers around talks and discussions to review knowledge on these topics and initiate the writing of perspective/opinion papers.

(3) Development of a collaborative project. The long-term objective of this transversal axis (within the next 4 years) is to develop a collaborative project that addresses some (or all) the issues/gaps identified in activities (1) and (2). We expect this project to integrate theoreticians and empiricists, and making use of the experimental facilities available at SETE and abroad.

Leaders

Jose Montoya and Staffan Jacob

Experimentation, observation, and causality

The use of experiments in ecology and evolution has increased dramatically in the last two or three decades, as attested by the recent construction of many highly-controlled experimental facilities and the surge of many long-term experiments in natural ecosystems. Although some reviews have already discussed the use of experimental facilities and the scientific insights that can be gained from them, there has been no comprehensive attempt to review the role of experiments in ecology, their constraints and limits, and the way to overcome these constraints and limits in the future (time, space, extreme environments, multiple stressors, new views on experimentation, combining experiments, observations and models, etc.).

SETE is well placed to launch a reflection on these issues. We therefore propose to organise a series of internal workshops (1-2 day workshops every six months) extended to some selected people from outside SETE. Here is a list of potential subjects to be addressed:

  • the use of model systems in ecological experiments and their limits;
  • reproducibility and the validity of ecological experiments;
  • statistics and causation;
  • feedbacks between ecological experiments, ecological theory and ecological modelling;
  • feedbacks between experiments and observations in natural systems;
  • experiments and scale: how long is long enough? How big is big enough?
  • evolutionary dynamics of complex ecosystems;
  • experimentation on feedbacks between ecosystem and humans;
  • experiments on mosaics of ecosystems;
  • frontiers in multiple stressors/factors experiments and the study of extreme events.

One objective of these discussions will be to produce a series of contributions, either as special issues or as independent papers.

Leaders

Jean Clobert

Eco-evolutionary feedbacks

Feedbacks between ecological and evolutionary dynamics are increasingly recognized as important. They are being studied at SETE using different model systems (protists, microbiota, butterflies, fish, crayfish, dragonflies) and a combination of approaches (observational, experimental, modelling) across teams. One focus is on dispersal, a key trait in an organism’s response to global changes, with strong effects on ecological and evolutionary dynamics. Dispersal evolves rapidly, shows genetic variation within metapopulations and modulates relationships between evolutionary and ecological parameters. Natural and anthropogenic habitat fragmentation affects evolution of dispersal, whose influence on population dynamics can cascade upwards to community dynamics and ecosystem functioning. At SETE, dispersal is studied by identifying and measuring multiple, potentially correlated traits that form heritable dispersal syndromes. This process enables modelling to make predictions that can be tested by experimental release of animals in microcosms and the terrestrial and aquatic metatrons.

A second focus of research on eco-evolutionary dynamics at SETE concerns intraspecific diversity.

Changes in intraspecific diversity have been demonstrated to generate evolutionary feedbacks acting on ecological dynamics, including ecosystem-level effects. For example, evolution of intraspecific functional diversity within crayfish populations influences the rate of litter decomposition in their lake habitat. Research at SETE also shows how ecological and evolutionary dynamics combine to influence processes as diverse as the assembly of the microbiome of sponges, the extinction probabilities of butterfly populations and the nestedness, diversity and resilience of mutualistic networks.

To animate this transversal axis, we plan to organize a couple of half-day workshops with presentations and discussions every year, notably around invited speakers in SETE’s seminar series who will talk on eco-evolutionary dynamics.

Leaders

Claire de Mazancourt and Michael Singer

Human-nature interactions

Globally increasing human population size, conversion of native lands to agriculture and urban sprawl have all led to a higher frequency and intensity of interactions between humans and nature. Research conducted at SETE has been used to inform issues that affect different sectors of society. These include conservation planning, improving ecosystem services by better understanding of feedbacks between human activities and ecosystem functioning, elucidating underlying psychology of social behaviour, and informing international policy on responding to changing biodiversity and changing climate. Unexpected consequences of increased human-nature interactions, such as transference of HIV virus from animal into human populations, can have devastating impacts. Although negative effects of intensive food production generate social pressures for sustainability, research at SETE has shown how time-delays in response to these pressures can, through their influence on ecosystem services, increase the vulnerability of social-ecological systems and render collapse of human populations more likely. SETE research has illuminated the importance of coordinating studies of human dependence on biodiversity with studies of human influences on biodiversity.

SETE is well-placed to delve deeper into these themes. One overarching question we hope to build upon is: what are the implications of increasing pressure from global change drivers on interactions between human populations and natural systems, and for feedback processes? Examples of the type of cross-team projects that could develop along the broad theme of human-nature interactions include:

(1) At the global scale, anthropogenic climate change and human transport of exotic plants and animals are driving movements of both wild species and of humans across the Earth. These shifts in geographic locations are spawning new interactions within both natural and human communities, and between humans and wild species. Emergent questions SETE researchers are beginning to address are: what are the current and future impacts of these novel interactions on local biodiversity, human health, food access and production, and human well-being? How can better understanding of the genetic vs plastic basis of cooperative behaviour, and its cultural and genetic evolution, help to inform policy actions aimed at altering public behaviour, e.g. to lower greenhouse gas emissions or protect biodiversity?

(2) At the regional scale, humans are increasingly encroaching into surrounding natural areas, creating fragmented and degraded habitats that are less resilient to either natural disturbance or increasing intensity of human perturbations. In turn, fragmented, degraded landscapes are less suitable for accommodating both wild species and humans that are being driven out of their native lands. How can better understanding of processes occurring during the formation of novel wild and human communities help to improve the resilience and stability of coupled human-natural systems?

Leader

Camille Parmesan

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CHANGE - Research Projects

CHANGE

Research Projects

GRACIL

Genome Rearrangements and Adaptation: response to salinity stress in a CILiate

Objectives

We aim at determining the role of genomic rearrangements in the adaptation to environmental changes. The project combines experimental evolution in ciliate microcosms to the generation of optical maps, a cutting-edge technic allowing to explore genome organization (inversions, deletions, duplications). We try to relate adaptive patterns in response to salinity stress to the target traits under selection (morphology, mobility) and the underlying genomic mechanisms. We hope to open new perspectives on the integration of under-explored molecular mechanisms in the outcome of evolution at contemporary time-scales.

Funding

FRAIB

Leaders

Delphine Legrand, Hervé Philippe and William Marande (CNRGV, INRA).

Participants:

Rick Verdonck

POLLUCLIM

Plasticity and adaptability to the combined effect of pollution and climate change

Objectives

The number and rate of anthropogenic alterations impose such intense selective pressures that biodiversity is irreversibly impacted. Plasticity and adaptability are key eco-evolutionary processes that could mitigate biodiversity loss. However, few studies have determined how the combined effects of anthropogenic stressors affect organisms’ immediate and evolutionary response.

POLLUCLIM will first determine the plastic response to warmer and/or polluted environments on a panel of ciliate genotypes in microcosms. Then, the probability of adaptation will be assessed, and we will determine if exposure to one stressor influences the response to another stressor. Finally, adaptive patterns will be related to genetic backgrounds and mutagenesis effects of stressors.

POLLUCLIM should improve our understanding of tolerance and adaptability patterns to multiple anthropogenic stressors, with access to the underlying molecular mechanisms.

Funding

ANR JCJC

Leader

Delphine Legrand

Participants:

Hervé Philippe, Michèle Huet, Staffan Jacob, Simon Blanchet

CHOOSE

Habitat choice: evolution and ecological consequences

Objectives

Growing evidence suggests that individuals that choose to leave a habitat are often phenotypically different from those that stay in this habitat. Moreover, individuals should benefit from settling in specific environments that maximize their fitness, which would create covariation between phenotypes and environmental conditions. The major consequences of non- random dispersal decisions that are both phenotype- and context- dependent have been recently developed under habitat choice theory.

Compared to random dispersal, this theory predicts that habitat choice based dispersal should generate spatial heterogeneity of phenotypes and thus lead to drastically different consequences for a variety of ecological and evolutionary dynamics such as range distribution, metapopulation dynamics and local adaptation. However, our comprehension of what drives the evolution of habitat choice and its ensuing consequences remains weak, especially because of the lack of experimental approaches dedicated to testing theoretical predictions. This research project thus aims at identifying the environmental drivers of habitat choice evolution, and quantify its consequences for ecological dynamics.

To do so, I will adopt an experimental approach using spatially explicit microcosms of an actively dispersing ciliate. This experimental system offers an excellent opportunity to validate theoretically-derived predictions and thus to provide breakthrough advances on the environmental drivers and consequences of dispersal evolution.

Funding

ANR JCJC

Leader

Staffan Jacob

Participants:

Delphine Legrand, Michèle Huet, Alexis Chaine, Jean Clobert, Julie Campana

PAPEL

Anthropic pressures and the role of protists in the regulation of lake eutrophication

Objectives

Human-induced alterations are profoundly modifying biodiversity. Mountain lakes constitute important water reservoirs with high patrimonial value.

They are however especially vulnerable to the accumulation of anthropic pressures, some of them showing signs of eutrophication. In this project, we aim at characterizing the role of protists in the eutrophication process of Pyrenean mountain lakes at several spatial and temporal scales.

Using an interdisciplinary approach merging metagenomics, metabarcoding, paleoecology (study of sediment cores), and protist microcosm experiments, we will try to reconstruct the dynamics and functioning of these ecosystems.

Leader

Delphine Legrand, Staffan Jacob

Participants:

Didier Galop (GEODE), Simon Blanchet, Hervé Philippe, Bart Haegeman, Michèle Huet, Jean Clobert, Morgane Gibert (AMIS), Ludovic Orlando (AMIS)

MICMAC

Exploring the interplay between mutualism and parasitism at micro- and macro-evolutionary scales

Objectives

The interactions between organisms affect gene and ecosystem diversity. Given that most species are constantly challenged by both mutualistic and parasitic microorganisms, we propose that mutualism and parasitism could influence each other.

Here, using plants as a model we will test the hypothesis that parasitism and mutualism impact the evolution of each other. To do this, we first propose (Partner 1) to identify genes involved in parasitism or mutualism using genetic approaches (GWAS, selection scans) and to compare their evolution within species of two deeply divergent clades of land plants, angiosperms and liverworts, including in species in which mutualism was lost (e.g., Arabidopsis thaliana and Marchantia polymorpha).

Then, through phylogenomic approaches across the entire embryophyte phylogeny (Partner 2), we will finely describe the evolution of these genes and infer selective processes to understand how selection can manage the interplay between mutualism and parasitism.

Leader

Pierre-Marc Delaux, Hervé Philippe

Participants:

Maxime Bonhomme

URBAN TIT

Urban environment on individual performance at a multi-populational level in great tits

Objectives

Effects of the Urbanization imposes important selection pressures on organisms, yet our understanding of how anthropogenic impacts often relies on small scale studies that focus on one population and one impact. In this project, we have created a network of labs within France to study anthropogenic impacts in a common songbird, the great tit, across 6 replicate urban-rural population pairs.

We will directly measure pollution at each site (heavy metals, pesticides) as well as overall fitness, characterize the microbiome, and look for genetic signatures of past selection associated with urbanization. In addition, we will conduct experimental studies to examine the impact of urbanization on cognitive performance in the wild and how light and sound pollution impact cognitive performance in captivity. As a whole, this study will provide a much larger scale assessment of how anthropogenic pollution impacts songbirds.

Funding

ANR 2019

Leader

S. Massemin is lead PI, U. Strasbourg; A. Chaine (SETE) and P. Heeb (Toulouse) are co-PIs with 5 other labs

Participants at Sete:

Maxime Cauchoix (Post-doc), Nory El Ksabi (IE), Maine Bely (IE)

SOCO

Causes and consequences of social competence and cognition:
from genes to fitness

Objectives

Cognitive abilities are critical to many social interactions yet few studies have examined the link between social plasticity (called social competence) and cognitive performance. In this study, we will examine the relationship between inter-individual variation in social plasticity and performance at specific cognitive tasks across an ecological gradient (1000m elevational gradient). In addition, we will examine how variation in cognition and social competence impact fitness in the wild and whether this effect is mediated through parental behavior.

Finally, we will examine the genetic basis of variation in cognitive performance and social competence. To achieve these goals, we will use a combination of experiments in the SETE aviaries as well as experiments in the field using a newly developed field cognitive testing device. Together, the elements of this study will provide a new perspective on how cognition and social plasticity interact as well as both the causes and consequences of inter-individual variation in those traits.

Funding

ANR 2018

Leaders

A. Chaine (SETE) ; Co-PIs : A. Charmantier (Montpellier), P. Heeb (Toulouse), N. Claidière (Aix en Provence)

Participants at SETE:

Nory El Ksabi (IE), Marine Bely (IE), Maxime Cauchoix (Post-doc), Thomas Crouchet (PhD student)

MICROTRAC

UHF microtracers, a new technology to track the movements and behaviors of micro wildlife

Objectives

To adapt to global warming and the constant increase in anthropogenic activity (pollution, urbanization, intensive agriculture, landscape fragmentation), wild animals must develop new behavioral strategies. Here, we focus on the birds of the sparrow group which are constantly decreasing and which play a key role in the biodiversity of the countryside, the prevention of agricultural pests and the functioning of ecosystems.

One of the key questions facing ecological researchers today is how animals adjust their movements to new changes in their environment linked to human activity. However, our understanding of the behavior of sparrows in their natural environment, and in particular in the face of anthropogenic activity, remains limited, as tools to track small sparrows do not currently exist.

In this project, we are collaborating with the company Xerius, (http://xerius.fr) which has unique expertise in radio frequency technology and animal telemetry to develop a new micro-plotter suitable for small passerines. With these new tracers, we will examine how urbanization influences movements, territoriality, and social interactions.

Funding

Région Occitanie Research and Society Grants (Public-Private partnerships in research)

Leaders

PIs: P. Heeb (Toulouse) is lead PI with Xerius Microsystems as the private partner; co-PIs : A. Chaine (SETE), A. Gregoire (Montpellier)

Participants at SETE:

Maxime Cauchoix (post-doc), Nory El Ksabi (IE)

Long term maintenance of striking color variation in Reunion Island Grey White-eyes:the role of social and reproductive behavior

Objectives

Variation among individuals within a species, especially in color traits, has attracted intense attention since the early explorers and naturalists brought back specimens from the far reaches of the world. Yet how variation is maintained within populations still presents a scientific puzzle since natural or social selection should reduce variation within populations unless there are some countervailing benefits. Reunion Island Grey White Eyes show a color polymorphism that has been stable for over 50 years and we are providing a new and more nuanced description of this color polymorphism and investigating the genetic basis of the color polymorphism as well as how selection acts on each morph through parasite resistance, reproductive success, and social interactions.

Funding

FRAIB Grant for collaborative research with C. Thébaud; National Geographic Research Grant

Leaders

PIs: A. Chaine at SEEM and C. Thebaud at Toulouse

Participants at SETE:

Maya Mould (PhD student)

Structure and resilience of social networks under population turnover

Objectives

All social networks change membership over time: individuals are lost through death or dispersal, and new individuals join the social network through birth and immigration. Yet we know almost nothing about how this universal process of population turnover shapes the structure of social networks, nor do we understand how social networks may or may not be resilient to population turnover. In this project, we are developing new theory on how turnover might impact social networks a confronting such models with long term empirical data on the social network of migrant golden-crowned sparrows (Zonotrichia atricapilla). Furthermore, we will use this long term data to understand how network stability impacts social behaviors, communication networks, and long term survival in this species.

Funding

NSF-CAREER

Leaders

D. Shizuka (U. Nebraska) as lead PI; co-PIs are A. Chaine (SETE) and B. Lyon (Univ Calif, Santa Cruz

ECONECT

Development of connected environmental sentinel systems to better understand the degradation of rivers, the decline of bees and birds

Objectives

The exponential development of human activity over the past two centuries has profoundly changed the environment in which we live in. In particular, urbanisation, industrialisation and intensive agriculture have generated, and still generate, various forms of pollution that play a major role in the loss of biodiversity and the global warming observed worldwide.

To carry out environmental policies of conservation and sustainable development, it is essential to understand by what mechanisms the pollution produced by human activity contributes to the degradation of ecosystems. This requires being able to both quantify the different types of pollution present and the responses of organisms to these environmental stresses in a fine and integrative manner. There are currently no tools to meet this challenge across an entire territory.

With Econect, we propose to develop new environmental sentinel systems and deploy them in the Occitanie region. These real field laboratories, autonomous, connected and scalable, will be able to both measure the pollution present in water, air or soil and the response to these stresses from bio-indicator organisms (freshwater algae, honey bee and great tit). The technological challenges and innovation related to this project are related to the development of new sensors, automatic and remote data recovery, and the management of a continuous and massive flow of data.

This project brings together manufacturers for the development of sentinel systems (Beeguard; Select Design) as well as data management (Adict Solutions). A network of 12 environmental sentinel systems will then be deployed in Occitanie according to three ecological gradients (altitude, urbanization and type of agriculture) characterised via a spatial analysis of land use and pollutants (Heavy metals, Pesticides). We will use a participatory science protocol (schools, associations, beekeepers, ornithologists, farmers) both for the local management of the equipment and to raise awareness of environmental issues in our region. For more information, check out : econect.cnrs.fr
 

Funding

Région Occitanie Recherche and FEDER

Leaders

Arnaud Elger (ECOLAB), Maxime Cauchoix (SETE)

Participants

Arnaud Elger (Toulouse), Alexis Chaine (SETE), Mathieu Lihoreau (CRCA), Rahim Kacimi (IRIT), Vincent Raimbault (LAAS), Marie-Pierre Julien (GEODE), entreprises Select Design, Adict Solutions, BeeGuard

PARTICIPANTS FROM SETE :

Maxime Cauchoix (researcher), Nory El Ksabi (engineer)

INTERREG POCTEFA ADAPYR

Capitalisation, observation, transfer and appropriation of climate change adaptation strategies in the Pyrenees in the context of cross-border cooperation

Objectives

OPCC ADAPYR is a unifying project of the Pyrenees in terms of observation, capitalization, transfer of knowledge and good practices towards resilience and  adaptation to climate change. Its objectives are to systematize the monitoring of climate impacts in the Pyrenees and to define a common Pyrenean adaptation strategy to climate change.

Along with the Communauté de Travail des Pyrénées, 12 beneficiary organizations and 30 partner organisations from all over the Pyrenees will bring and share their knowledge and work on the impacts, vulnerability and adaptation to climate change in various fields such as flora, fauna, lakes and peat bogs, glaciers, forests, forests, watersheds, natural hazards or the study of climate. The partner SETE, represented by Fabien Aubret, studies within the framework of ADAPYR the effects of climate change on the ecology of a lizard species endemic to the Pyrenees, the Bonnal lizard). Iberolacerta lizards are all endemic to the Pyrenees, registered on the IUCN world red list (2015) and benefit from a National Action Plan supported by Nature Midi-Pyrénées (NMP). 

The study will make it possible to identify the populations of Iberolacerta bonnali on the Pyrenean chain, and to propose sustainable development strategies in the face of climate change in order to limit the loss of biodiversity.

The study will sample populations of Iberolacerta bonnali, experimentally assess their physiological adaptations to high life and propose sustainable development strategies in the face of climate change in order to limit the loss of biodiversity.

THe OPCC ADAPYR Website :

https://www.opcc-ctp.org/es/proyecto/opcc-adapyr

https://www.opcc-ctp.org/fr/proyecto/opcc-adapyr

Funding

FEDER, European Union  

Leaders

Communauté de Travail des Pyrénées ; Olivier Guillaume, Fabien Aubret, Laura Kouyoumdjian (IE)

Urban Wetlands Conservation

Objectives

Impact of urbanization on tiger snakes in the wetlands of Perth and surrounding areas. This research project (Doctoral Thesis) measures whether and how environmental degradation by urbanization and pollution makes vertebrates in wetlands more susceptible to diseases and parasites, by comparing the health status and ecology populations of tiger snakes along geographic and historical gradients.

 

This project assesses the degradation of wetland health (habitat structure and water quality) through an urban matrix to determine the effects of bioaccumulation of contaminants in tiger snakes, the effects of degradation of wetlands on parasitism in tiger snakes and integrate all information to determine whether tiger snakes can be used as an ecological indicator of wetland health.

Funding

Curtin University

Team

Pr Bill Bateman (Curtin University, Perth, Australie), Fabien Aubret (SETE), Monique Gagnon (Curtin University, Perth, Australie), Damian Lettoof (PhD student).

AQUATHERM

The potential of hydroregulation and thermoregulation to influence ecological responses to climate change

Objectives

The secular changes in environmental temperatures and water availability driven by climate change affect the physiological performances of ectothermic animals and push some of their populations on a fast lane to extinction. The sensitivity, resilience and adaptive potential to climate change of ectotherms are all largely determined by physiological and behavioural capacities and tolerances. Individual responses to changes in temperature and water availability involve thermoregulation (i.e., physiological and behavioural regulation of body temperature) and hydroregulation (i.e., physiological and behavioural regulation of the water balance). In ectotherms, responses to water constraints remain poorly investigated when considering climate change.

Yet, it is anticipated that hydroregulation and thermoregulation will influence each other through complex, possibly conflicting pathways leading to ecological responses to climate change difficult to predict by focusing solely on thermal biology. How this interplay between thermoregulation and hydroregulation influences vulnerability to climate change remains largely unknown because we lack studies that examine jointly hydro- and thermoregulation strategies involved in response to climate change. One promising and comprehensive approach to tackle this problem is to use heat, mass and water budget models that are robust and sufficiently general to be applied to a large range of study systems.

Here, we will use ecophysiology and behavioural ecology to enhance our understanding of this critical facet in terrestrial ectotherms. Focusing on squamate reptiles (lizards and snakes), we will combine mechanistic biophysical models, empirical studies of physiological and behavioural traits at the individual and population levels using two model species from two French Mountain ranges, climate niche simulations for these species, and comparative analyses across all squamate reptiles. We will be able to describe and understand for the first time the covariation patterns between thermoregulation and hydroregulation, and to investigate and improve our capacity to predict ecological effects of two global change pressures (temperature and water) in terrestrial ectotherms.

This approach will provide new insights on the role of proximate functional traits in determining species distribution and sensitivity to climate change, and translate into knowledge applicable in other terrestrial ectotherms and wildlife management.

Funding

ANR

Leaders

Jean-François Le Galliard, Sorbonne University, Paris, France, Fabien Aubret (SETE)

Origin, maintenance and function of color polymorphisms

Objectives

One of the main goal of this project is the identification of mutations and genes associated with specific phenotypes. In this case, the information concerns genes involved in vertebrate coloration, which at first glance may appear to have no bearing on the study of human health. However, any knowledge that clarifies the role of individual genes in complex metabolic cycles, especially in vertebrates, contributes to a better understanding of the human genome and may eventually have an application in preventing disease in humans and other animals. By way of example, we cite a gene involved in Drosophila pigmentation (SPR, sepia pterin reductase), which may possibly be involved in the orange coloration of lizards under study in this project, which is associated with serious human diseases (https://ghr.nlm.nih.gov/gene/SPR#conditions ). There may be other genes associated with pigmentation whose knowledge proves to be important in the study of diseases in humans, and all the knowledge that can be gained about their function (especially in the context of pigmentation, which is often part of metabolic pathways) will certainly be useful in the study of human disease.

Funding

Portugal 2020, European Union

leaders

Dr Catarina Pinho, CIBIO, University of Porto, Portugal, Fabien Aubret (SETE)

The molecular basis of adaptation to environmental change: tiger snake epigenetics

Objectives

This project aims to investigate mechanisms underlying animal adaptation to future environmental change by examining the molecular basis for phenotypic plasticity in snakes. This project will specifically examine variation in genetic/epigenetic profiles and compare against important fitness traits, such as variable head size, bite force and swallowing performance to identify relationships between molecular change and physiology. Such research is a critical first step in improving our knowledge of the mechanisms whereby animal populations may adapt to environmental change, allowing us to facilitate such processes or concentrate conservation effort where species are unable to adapt via epigenetic modification.

Leaders

Dr Vicki Thomson, University of Adelaide, Australia, Fabien Aubret (SETE)

Mention légales

Mentions légales

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