Model-checking ecological state-transition graphs.

Model-checking is a methodology developed in computer science to automatically assess the dynamics of discrete systems, by checking if a system modelled as a state-transition graph satisfies a dynamical property written as a temporal logic formula. The dynamics of ecosystems have been drawn as state-transition graphs for more than a century, ranging from state-and-transition models to assembly graphs. Model-checking can provide insights into both empirical data and theoretical models, as long as they sum up into state-transition graphs. While model-checking proved to be a valuable tool in systems biology, it remains largely underused in ecology apart from precursory applications. This article proposes to address this situation, through an inventory of existing ecological STGs and an accessible presentation of the model-checking methodology. This overview is illustrated by the application of model-checking to assess the dynamics of a vegetation pathways model. We select management scenarios by model-checking Computation Tree Logic formulas representing management goals and built from a proposed catalogue of patterns. In discussion, we sketch bridges between existing studies in ecology and available model-checking frameworks. In addition to the automated analysis of ecological state-transition graphs, we believe that defining ecological concepts with temporal logics could help clarify and compare them.

Environmental spillover of emerging viruses: Is it true?

The concept of environmental "spillover" of pathogens to humans is widely used in the scientific literature about emerging diseases with the idea that it is scientifically proven. However, the exact characterization of the mechanism of spillover is simply lacking. A systematic review retrieved 688 articles using this term. The systematic analysis revealed an irreducible polysemy covering ten different definitions. It also demonstrated the absence of explicit definition in most of the articles, and even antinomies. A modeling analysis of the various processes described by these ten definitions showed that none of them corresponded to the complete trajectory leading to the emergence of a disease. There is no article demonstrating a mechanism of spillover. There are only ten articles proposing ideas on how a putative spillover could work but they merely are intellectual constructions. All other articles only reuse the term with no demonstration. It is essential to understand that since there is no scientific concept behind the "spillover", it might be dangerous to base public health and public protection against future pandemics on it.

On the History of Ecosystem Dynamical Modeling: The Rise and Promises of Qualitative Models.

Ecosystem modeling is a complex and multidisciplinary modeling problem which emerged in the 1950s. It takes advantage of the computational turn in sciences to better understand anthropogenic impacts and improve ecosystem management. For that purpose, ecosystem simulation models based on difference or differential equations were built. These models were relevant for studying dynamical phenomena and still are. However, they face important limitations in data-poor situations. As a response, several formal and non-formal qualitative dynamical modeling approaches were independently developed to overcome some limitations of the existing methods. Qualitative approaches allow studying qualitative dynamics as relevant abstractions of those provided by quantitative models (e.g., response to press perturbations). Each modeling framework can be viewed as a different assemblage of properties (e.g., determinism, stochasticity or synchronous update of variable values) designed to satisfy some scientific objectives. Based on four stated objectives commonly found in complex environmental sciences ((1) grasping qualitative dynamics, (2) making as few assumptions as possible about parameter values, (3) being explanatory and (4) being predictive), our objectives were guided by the wish to model complex and multidisciplinary issues commonly found in ecosystem modeling. We then discussed the relevance of existing modeling approaches and proposed the ecological discrete-event networks (EDEN) modeling framework for this purpose. The EDEN models propose a qualitative, discrete-event, partially synchronous and possibilistic view of ecosystem dynamics. We discussed each of these properties through ecological examples and existing analysis techniques for such models and showed how relevant they are for environmental science studies.

How climate, migration ability and habitat fragmentation affect the projected future distribution of European beech.

Recent efforts to incorporate migration processes into species distribution models (SDMs) are allowing assessments of whether species are likely to be able to track their future climate optimum and the possible causes of failing to do so. Here, we projected the range shift of European beech over the 21st century using a process-based SDM coupled to a phenomenological migration model accounting for population dynamics, according to two climate change scenarios and one land use change scenario. Our model predicts that the climatically suitable habitat for European beech will shift north-eastward and upward mainly because (i) higher temperature and precipitation, at the northern range margins, will increase survival and fruit maturation success, while (ii) lower precipitations and higher winter temperature, at the southern range margins, will increase drought mortality and prevent bud dormancy breaking. Beech colonization rate of newly climatically suitable habitats in 2100 is projected to be very low (1-2% of the newly suitable habitats colonised). Unexpectedly, the projected realized contraction rate was higher than the projected potential contraction rate. As a result, the realized distribution of beech is projected to strongly contract by 2100 (by 36-61%) mainly due to a substantial increase in climate variability after 2050, which generates local extinctions, even at the core of the distribution, the frequency of which prevents beech recolonization during more favourable years. Although European beech will be able to persist in some parts of the trailing edge of its distribution, the combined effects of climate and land use changes, limited migration ability, and a slow life-history are likely to increase its threat status in the near future.

Origins of evolution: non-acquired characters dominates over acquired characters in changing environment.

Natural Selection is so ubiquitous that we never wonder how it appeared as the evolution rule driving Life. We usually wonder how Life appeared, and seldom do we make an explicit distinction between Life and natural selection. Here, we apply the evolution concept commonly used for studying Life to evolution itself. More precisely, we developed two models aiming at selecting among different evolution rules competing for their supremacy. We explored competition between acquired (AQ) versus non-acquired (NAQ) character inheritance. The first model is parsimonious and non-spatial, in order to understand relationships between environmental forcings and rule selection. The second model is spatially explicit and studies the adaptation differences between AQ and NAQ populations. We established that NAQ evolution rule is dominating in case of changing environment. Furthermore, we observed that a more adapted population better fits its environmental constraints, but fails in rapidly changing environments. NAQ principle and less adapted populations indeed act as a reservoir of traits that helps populations to survive in rapidly changing environments, such as the ones that probably Life experienced at its origins. Although perfectible, our modeling approaches will certainly help us to improve our understanding of origins of Life and Evolution, on Earth or elsewhere.

Dominance and species co-occurrence in highly diverse ant communities: a test of the interstitial hypothesis and discovery of a three-tiered competition cascade.

The role of competitive exclusion is problematic in highly diverse ant communities where exceptional species richness occurs in the face of exceptionally high levels of behavioural dominance. A possible non-niche-based explanation is that the abundance of behaviourally dominant ants is highly patchy at fine spatial scales, and subordinate species act as insinuators by preferentially occupying these gaps--we refer to this as the interstitial hypothesis. To test this hypothesis, we examined fine-scale patterns of ant abundance and richness according to a three-tiered competition hierarchy (dominants, subdominants and subordinates) in an Australian tropical savanna using pitfall traps spaced at 2 m intervals. Despite the presence of gaps in the fine-scale abundance of individual species, the combined abundance of dominant ants (species of Iridomyrmex, Papyirus and Oecophylla) was relatively uniform. There was therefore little or no opportunity for subordinate species to preferentially occupy gaps in the foraging ranges of dominant species, and we found no relationship between the abundance of dominant ants and nondominant species richness at fine spatial scales. However, we found a negative relationship between subdominant and subordinate ants, a negative relationship between dominant and subdominant ants, and a positive relationship between dominant and subordinate ants. These results suggest that dominant species actually promote species richness by neutralizing the effects of subdominant species on subordinate species. Such indirect interactions have very close parallels with three-tiered trophic cascades in food webs, and we propose a "competition cascade" where the interactions are through a competition rather than trophic hierarchy.

[Strengths, weaknesses, and opportunities of French research in trophic ecology]. / Forces, failles et opportunités de la recherche française en écologie trophique.

The French National Institute of Ecology and Environment (INEE) aims at fostering pluridisciplinarity in Environmental Science and, for that purpose, funds ex muros research groups (GDR) on thematic topics. Trophic ecology has been identified as a scientific field in ecology that would greatly benefit from such networking activity, as being profoundly scattered. This has motivated the seeding of a GDR, entitled "GRET". The contours of the GRET's action, and its ability to fill these gaps within trophic ecology at the French national scale, will depend on the causes of this relative scattering. This study relied on a nationally broadcasted poll aiming at characterizing the field of trophic ecology in France. Amongst all the unique individuals that fulfilled the poll, over 300 belonged at least partly to the field of trophic ecology. The sample included all French public research institutes and career stages. Three main disruptions within the community of scientist in trophic ecology were identified. The first highlighted the lack of interfaces between microbial and trophic ecology. The second evidenced that research questions were strongly linked to single study fields or ecosystem type. Last, research activities are still quite restricted to the ecosystem boundaries. All three rupture points limit the conceptual and applied progression in the field of trophic ecology. Here we show that most of the disruptions within French Trophic Ecology are culturally inherited, rather than motivated by scientific reasons or justified by socio-economic stakes. Comparison with the current literature confirms that these disruptions are not necessarily typical of the French research landscape, but instead echo the general weaknesses of the international research in ecology. Thereby, communication and networking actions within and toward the community of trophic ecologists, as planned within the GRET's objectives, should contribute to fill these gaps, by reintegrating microbes within trophic concepts and setting the seeds for trans- and meta-ecosystemic research opportunities. Once the community of trophic ecologists is aware of the scientific benefit in pushing its boundaries forwards, turning words and good intentions into concrete research projects will depend on the opportunities to obtain research funding.

Language: A fresh concept to integrate syntactic and semantic information in life sciences.

Several fields in biology tend to view the concept of information from one or the other of two extreme positions. Exclusionists base their stance of total rejection on gene-centrism and gene-determinism, typified by the recently-established endo-Darwinist school of life sciences. At the other end of the spectrum, there is total acceptance, as in the newly developed information-centred paradigms that populate biosemiotics. We propose in this paper to split the informational concepts into two irreducible (but linked) poles: the syntactic (concerned with the quantification of the information structure or complexity in a system), and the semantic (concerned with the organization rules and causality weights of interactions in a system). We claim that the past and present uses of the concept could then be classified as various degrees of oscillation between the two poles. The concept of language presents itself as a good tool with which to bridge the syntactic and the semantic poles, combining as it does the form-related and the meaning-related aspects of information, while methodologically supporting formal grammatical models in life sciences. We aim to show, at the same time, that neither of these poles alone can suffice to efficiently and holistically describe, model, and predict natural phenomena.

Honey pollen: using melissopalynology to understand foraging preferences of bees in tropical South India.

The aim of the study was to use melissopalynology to delineate the foraging preferences of bees in tropical environs. This was done by comparing pollen spectra obtained from the same hives every three months for three years at four sampling locations (in two sites) within a confined landscape mosaic. If melissopalynology is highly replicable, the spatial variation of the pollen spectrum from the honey samples would be much more than the temporal (inter-annual) variations. In other words, given the three factors, Month, Year and Location, honey pollen from different Locations, in a given Year and Month, would be much less similar than samples from different Years, in a given Location and Month. We then determined how the factors, Month, Year and Location, influenced the pollen influx of honey. The pollen analyses of the 42 honey samples collected during the three years yielded 80 pollen taxa/types: 72 dicotyledonous and 8 monocotyledonous, encompassing 41 botanical families spread into seven life forms namely, trees, shrubs, epiphytes, herbs, climbers, grasses, and sedges. Our results showed that pollen spectra were equally comparable between Locations and between Months and Years; the importance of this result is that it helped to demonstrate the complexity of ecological/environmental phenomena involved in the process of foraging by bees in a heterogeneous and complex landscape.

Overview of the Distribution, Habitat Association and Impact of Exotic Ants on Native Ant Communities in New Caledonia.

Ants are among the most ubiquitous and harmful invaders worldwide, but there are few regional studies of their relationships with habitat and native ant communities. New Caledonia has a unique and diverse ant fauna that is threatened by exotic ants, but broad-scale patterns of exotic and native ant community composition in relation to habitat remain poorly documented. We conducted a systematic baiting survey of 56 sites representing the main New Caledonian habitat types: rainforest on ultramafic soils (15 sites), rainforest on volcano-sedimentary soils (13), maquis shrubland (15), Melaleuca-dominated savannas (11) and Acacia spirorbis thickets (2). We collected a total of 49 species, 13 of which were exotic. Only five sites were free of exotic species, and these were all rainforest. The five most abundant exotic species differed in their habitat association, with Pheidole megacephala associated with rainforests, Brachymyrmex cf. obscurior with savanna, and Wasmannia auropunctata and Nylanderia vaga present in most habitats. Anoplolepis gracilipes occurred primarily in maquis-shrubland, which contrasts with its rainforest affinity elsewhere. Multivariate analysis of overall ant species composition showed strong differentiation of sites according to the distribution of exotic species, and these patterns were maintained at the genus and functional group levels. Native ant composition differed at invaded versus uninvaded rainforest sites, in the absence of differences in habitat variables. Generalised Myrmicinae and Forest Opportunists were particularly affected by invasion. There was a strong negative relationship between the abundance of W. auropunctata and native ant abundance and richness. This emphasizes that, in addition to dominating many ant communities numerically, some exotic species, and in particular W. auropunctata, have a marked impact on native ant communities.

How can model comparison help improving species distribution models?

Today, more than ever, robust projections of potential species range shifts are needed to anticipate and mitigate the impacts of climate change on biodiversity and ecosystem services. Such projections are so far provided almost exclusively by correlative species distribution models (correlative SDMs). However, concerns regarding the reliability of their predictive power are growing and several authors call for the development of process-based SDMs. Still, each of these methods presents strengths and weakness which have to be estimated if they are to be reliably used by decision makers. In this study we compare projections of three different SDMs (STASH, LPJ and PHENOFIT) that lie in the continuum between correlative models and process-based models for the current distribution of three major European tree species, Fagussylvatica L., Quercusrobur L. and Pinussylvestris L. We compare the consistency of the model simulations using an innovative comparison map profile method, integrating local and multi-scale comparisons. The three models simulate relatively accurately the current distribution of the three species. The process-based model performs almost as well as the correlative model, although parameters of the former are not fitted to the observed species distributions. According to our simulations, species range limits are triggered, at the European scale, by establishment and survival through processes primarily related to phenology and resistance to abiotic stress rather than to growth efficiency. The accuracy of projections of the hybrid and process-based model could however be improved by integrating a more realistic representation of the species resistance to water stress for instance, advocating for pursuing efforts to understand and formulate explicitly the impact of climatic conditions and variations on these processes.

Understanding patchy landscape dynamics: towards a landscape language.

Patchy landscapes driven by human decisions and/or natural forces are still a challenge to be understood and modelled. No attempt has been made up to now to describe them by a coherent framework and to formalize landscape changing rules. Overcoming this lacuna was our first objective here, and this was largely based on the notion of Rewriting Systems, also called Formal Grammars. We used complicated scenarios of agricultural dynamics to model landscapes and to write their corresponding driving rule equations. Our second objective was to illustrate the relevance of this landscape language concept for landscape modelling through various grassland managements, with the final aim to assess their respective impacts on biological conservation. For this purpose, we made the assumptions that a higher grassland appearance frequency and higher land cover connectivity are favourable to species conservation. Ecological results revealed that dairy and beef livestock production systems are more favourable to wild species than is hog farming, although in different ways. Methodological results allowed us to efficiently model and formalize these landscape dynamics. This study demonstrates the applicability of the Rewriting System framework to the modelling of agricultural landscapes and, hopefully, to other patchy landscapes. The newly defined grammar is able to explain changes that are neither necessarily local nor Markovian, and opens a way to analytical modelling of landscape dynamics.