Unité Évolution, Écologie et Paléontologie

Summary In a rapidly changing world, ecology has the potential to move from empirical and conceptual stages to application and management issues. It is now possible to make large‐scale predictions up to continental or global scales, ranging from the future distribution of biological diversity to changes in ecosystem functioning and services. With these recent developments, ecology has a historical opportunity to become a major actor in the development of a sustainable human society. With this opportunity, however, also comes an important responsibility in developing appropriate predictive models, correctly interpreting their outcomes and communicating their limitations. There is also a danger that predictions grow faster than our understanding of ecological systems, resulting in a gap between the scientists generating the predictions and stakeholders using them (conservation biologists, environmental managers, journalists, policymakers). Here, we use the context provided by the current surge of ecological predictions on the future of biodiversity to clarify what prediction means, and to pinpoint the challenges that should be addressed in order to improve predictive ecological models and the way they are understood and used. Synthesis and applications . Ecologists face several challenges to ensure the healthy development of an operational predictive ecological science: (i) clarity on the distinction between explanatory and anticipatory predictions; (ii) developing new theories at the interface between explanatory and anticipatory predictions; (iii) open data to test and validate predictions; (iv) making predictions operational; and (v) developing a genuine ethics of prediction.

The diversity of life and its organization in networks of interacting species has been a long-standing theoretical puzzle for ecologists. Ever since May's provocative paper challenging whether 'large complex systems [are] stable' various hypotheses have been proposed to explain when stability should be the rule, not the exception. Spatial dynamics may be stabilizing and thus explain high community diversity, yet existing theory on spatial stabilization is limited, preventing comparisons of the role of dispersal relative to species interactions. Here we incorporate dispersal of organisms and material into stability-complexity theory. We find that stability criteria from classic theory are relaxed in direct proportion to the number of ecologically distinct patches in the meta-ecosystem. Further, we find the stabilizing effect of dispersal is maximal at intermediate intensity. Our results highlight how biodiversity can be vulnerable to factors, such as landscape fragmentation and habitat loss, that isolate local communities.

The Ordovician biodiversification has been recognized since the 1960s; the term 'The Great Ordovician Biodiversification Event', abbreviated by many as the 'GOBE', has been used for the past 20 years. The conceptual development and terminology applied to this crucial episode in marine life signify its considerable complexity. The GOBE includes successive biodiversity phases of the pelagic and benthic biotas, possibly decoupled. Put simply, the GOBE can be seen as a sequence of diversifications of the planktonic (late Cambrian-Early Ordovician), level-bottom benthic (Early-Middle Ordovician) and reef communities (Middle-Late Ordovician), although the boundaries of these 'events' are diachronous (as for the entire GOBE), and it is logical to assume that these communities co-evolved and interacted. The GOBE also includes several Biotic Immigration Events (BIMEs), such as the 'Richmondian Invasion' and the 'Boda Event', recording the large-scale dispersal of taxa from one biogeographical area to another. The GOBE is thus the sum of the diversity trends of all individual fossil groups showing rapid increases, diachronously, during different intervals and across different regions. It thus spans the entire Ordovician, capturing the increasing total diversity of marine organisms during the period. The GOBE is not simply one, but many sequential events.

The evolutionary transition from a solitary to a social lifestyle entails an elevated parasite cost because the social proximity associated with group living favors parasite transmission. Despite this cost, sociality is widespread in a large range of taxonomic groups. In this context, hosts would be expected to have evolved behavioral mechanisms to reduce the risk of parasite infection. Few empirical studies have focused on the influence of pathogen-mediated selection on the evolution of antiparasitic behavior in wild vertebrates. We report an adaptive functional relationship between parasitism and social behavior in mandrills, associated with evidence that they are able to gauge parasite status of their group members. Using long-term observations, controlled experiments, and chemical analyses, we show that (i) wild mandrills avoid grooming conspecifics infected with orofecally transmitted parasites; (ii) mandrills receive significantly more grooming after treatment that targets these parasites; (iii) parasitism influences the host's fecal odors; and (iv) mandrills selectively avoid fecal material from parasitized conspecifics. These behavioral adaptations reveal that selecting safe social partners may help primates to cope with parasite-mediated costs of sociality and that "behavioral immunity" plays a crucial role in the coevolutionary dynamics between hosts and their parasites.

The genomes of most plant species are dominated by transposable elements (TEs). Once considered as 'junk DNA', TEs are now known to have a major role in driving genome evolution. Over the last decade, it has become apparent that some stress conditions and other environmental stimuli can drive bursts of activity of certain TE families and consequently new TE insertions. These can give rise to altered gene expression patterns and phenotypes, with new TE insertions sometimes causing flanking genes to become transcriptionally responsive to the same stress conditions that activated the TE in the first place. Such connections between TE-mediated increases in diversity and an accelerated rate of genome evolution provide powerful mechanisms for plants to adapt more rapidly to new environmental conditions. This review will focus on environmentally induced transposition, the mechanisms by which it alters gene expression, and the consequences for plant genome evolution and breeding.

The Early Palaeozoic phytoplankton (acritarch) radiation paralleled a long-term increase in sea level between the Early Cambrian and the Late Ordovician. In the Late Cambrian, after the SPICE delta(13)C(carb) excursion, acritarchs underwent a major change in morphological disparity and their taxonomical diversity increased to reach highest values during the Middle Ordovician (Darriwilian). This highest phytoplankton diversity of the Palaeozoic was possibly the result of palaeogeography (greatest continental dispersal) and major orogenic and volcanic activity, which provided maximum ecospace and large amounts of nutrients. With its warm climate and high atmospheric CO(2) levels, the Ordovician was similar to the Cretaceous: a period when phytoplankton diversity was at its maximum during the Mesozoic. With increased phytoplankton availability in the Late Cambrian and Ordovician a radiation of zooplanktonic organisms took place at the same time as a major diversification of suspension feeders. In addition, planktotrophy originated in invertebrate larvae during the Late Cambrian-Early Ordovician. These important changes in the trophic chain can be considered as a major palaeoecological revolution (part of the rise of the Palaeozoic Evolutionary Fauna of Sepkoski). There is now sufficient evidence that this trophic chain revolution was related to the diversification of the phytoplankton, of which the organic-walled fraction is partly preserved.

Klug, C., Kroger, B., Kiessling, W., Mullins, G.L., Servais, T., Frýda, J., Korn, D. & Turner, S. 2009: The Devonian nekton revolution. Lethaia, 10.1111/j.1502-3931.2009.00206.x Traditional analyses of Early Phanerozoic marine diversity at the genus level show an explosive radiation of marine life until the Late Ordovician, followed by a phase of erratic decline continuing until the end of the Palaeozoic, whereas a more recent analysis extends the duration of this early radiation into the Devonian. This catch-all approach hides an evolutionary and ecological key event long after the Ordovician radiation: the rapid occupation of the free water column by animals during the Devonian. Here, we explore the timing of the occupation of the water column in the Palaeozoic and test the hypothesis that ecological escalation led to fundamental evolutionary changes in the mid-Palaeozoic marine water column. According to our analyses, demersal and nektonic modes of life were probably initially driven by competition in the diversity-saturated benthic habitats together with the availability of abundant planktonic food. Escalatory feedback then promoted the rapid rise of nekton in the Devonian as suggested by the sequence and tempo of water-column occupation. □Devonian, diversity, ecology, food webs, nekton, plankton, radiation.

The recent emergence of powerful genomic tools, such as high‐throughput genomics, transcriptomics and metabolomics, combined with the study of gnotobiotic animals, have revealed overwhelming impacts of gut microbiota on the host phenotype. In addition to provide their host with metabolic functions that are not encoded in its own genome, evidence is accumulating that gut symbionts affect host traits previously thought to be solely under host genetic control, such as development and behavior. Metagenomics and metatranscriptomics studies further revealed that gut microbial communities can rapidly respond to changes in host diet or environmental conditions through changes in their structural and functional profiles, thus representing an important source of metabolic flexibility and phenotypic plasticity for the host. Hence, gut microbes appear to be an important factor affecting host ecology and evolution which is, however, not accounted for in life‐history theory, or in classic population genetics, ecological and eco‐evolutionary models. In this forum, we shed new light on life history and eco‐evolutionary dynamics by viewing these processes through the lens of host– microbiota interactions. We follow a three‐level approach. First, current knowledge on the role of gut microbiota in host physiology and behavior points out that gut symbionts can be a crucial medium of life‐history strategies. Second, the particularity of the microbiota is based on its multilayered structure, composed of both a core microbiota, under host genetic and immune control, and a flexible pool of microbes modulated by the environment, which differ in constraints on their maintenance and in their contribution to host adaptation. Finally, gut symbionts can drive the ecological and evolutionary dynamics of their host through effects on individual, population, community and ecosystem levels. In conclusion, we highlight some future perspectives for integrative studies to test hypotheses on life history and eco‐evolutionary dynamics in light of the gut microbiota.

Extreme marine environments cover more than 50% of the Earth's surface and offer many opportunities for investigating the biological responses and adaptations of organisms to stressful life conditions. Extreme marine environments are sometimes associated with ephemeral and unstable ecosystems, but can host abundant, often endemic and welladapted meiofaunal species. In this review, we present an integrated view of the biodiversity, ecology and physiological responses of marine meiofauna inhabiting several extreme marine environments (mangroves, submarine caves, Polar ecosystems, hypersaline areas, hypoxic/anoxic environments, hydrothermal vents, cold seeps, carcasses/sunken woods, deep-sea canyons, deep hypersaline anoxic basins [DHABs] and hadal zones). Foraminiferans, nematodes and copepods are abundant in almost all of these habitats and are dominant in deep-sea ecosystems. The presence and dominance of some other taxa that are normally less common may be typical of certain extreme conditions. Kinorhynchs are particularly well adapted to cold seeps and other environments that experience drastic changes in salinity, rotifers are well represented in polar ecosystems and loriciferans seem to be the only metazoan able to survive multiple stressors in DHABs. As well as natural processes, human activities may generate stressful conditions, including deoxygenation, acidification and rises in

Understanding the mechanisms responsible for stability and persistence of ecosystems is one of the greatest challenges in ecology. Robert May showed that, contrary to intuition, complex randomly built ecosystems are less likely to be stable than simpler ones. Few attempts have been tried to test May's prediction empirically, and we still ignore what is the actual complexity-stability relationship in natural ecosystems. Here we perform a stability analysis of 116 quantitative food webs sampled worldwide. We find that classic descriptors of complexity (species richness, connectance and interaction strength) are not associated with stability in empirical food webs. Further analysis reveals that a correlation between the effects of predators on prey and those of prey on predators, combined with a high frequency of weak interactions, stabilize food web dynamics relative to the random expectation. We conclude that empirical food webs have several non-random properties contributing to the absence of a complexity-stability relationship.

Selection on codon usage bias is well documented in a number of microorganisms. Whether codon usage is also generally shaped by natural selection in large organisms, despite their relatively small effective population size (Ne), is unclear. In animals, the population genetics of codon usage bias has only been studied in a handful of model organisms so far, and can be affected by confounding, nonadaptive processes such as GC-biased gene conversion and experimental artefacts. Using population transcriptomics data, we analyzed the relationship between codon usage, gene expression, allele frequency distribution, and recombination rate in 30 nonmodel species of animals, each from a different family, covering a wide range of effective population sizes. We disentangled the effects of translational selection and GC-biased gene conversion on codon usage by separately analyzing GC-conservative and GC-changing mutations. We report evidence for effective translational selection on codon usage in large-Ne species of animals, but not in small-Ne ones, in agreement with the nearly neutral theory of molecular evolution. C- and T-ending codons tend to be preferred over synonymous G- and A-ending ones, for reasons that remain to be determined. In contrast, we uncovered a conspicuous effect of GC-biased gene conversion, which is widespread in animals and the main force determining the fate of AT↔GC mutations. Intriguingly, the strength of its effect was uncorrelated with Ne.

The hepatitis C virus (HCV) genome encodes two envelope glycoproteins (E1 and E2). These glycoproteins interact to form a noncovalent heterodimeric complex which in the cell accumulates in endoplasmic reticulum (ER)-like structures. The transmembrane domain of E2, at least, is involved in HCV glycoprotein complex localization in this compartment. In principle, ER localization of a protein can be the consequence of actual retention in this organelle or of retrieval from the Golgi. To determine which of these two mechanisms is responsible for HCV glycoprotein complex accumulation in the ER, the precise localization of these proteins was studied by immunofluorescence, and the processing of their glycans was analyzed. Immunolocalization of HCV glycoproteins after nocodazole treatment suggested an ER retention. In addition, HCV glycoprotein glycans were not modified by Golgi enzymes, indicating that the ER localization of these proteins is not because of their retrieval from the cis Golgi. Retention of HCV glycoprotein complexes in the ER without retrieval suggests that this compartment plays an important role for the acquisition of the envelope of HCV particles. A true retention in the ER was also observed for E2 expressed in the absence of E1 or for a chimeric protein containing the ectodomain of CD4 in fusion with the transmembrane domain of E2. These data indicate that, in HCV glycoprotein complex, the transmembrane domain of E2, at least, is responsible for true retention in the ER, without recycling through the Golgi.

ABSTRACT Aim Africa is identified by the Inter‐governmental Panel on Climate Change (IPCC) as the least studied continent in terms of ecosystem dynamics and climate variability. The aim of this study was (1) to adapt the Lund‐Postdam‐Jena‐GUESS (LPJ‐GUESS) ecological modelling framework to Africa by providing new parameter values for tropical plant functional types (PFT), and (2) to assess the sensitivity of some African biomes to changes in precipitation regime. Location The study area was a representative transect (0–22° N and 7–18° E) through the transition from equatorial evergreen forests to savannas, steppes and desert northwards. The transect showed large latitudinal variation in precipitation (mean rainfall ranged from 50 to 2300 mm year −1 ). Methods New PFT parameters used to calibrate LPJ‐GUESS were based on modern pollen PFTs and remote sensed leaf area index (LAI). The model was validated using independent modern pollen assemblages, LAI and through comparison with White's modern potential vegetation map. Several scenarios were developed by combining changes in total rainfall amount with variation in the length of the dry season in order to test the sensitivity of African biomes. Results Simulated vegetation compared well to observed data at local and regional scales, in terms of ecosystem functioning (LAI), and composition (pollen and White's vegetation map). The assessment of the sensitivity of biomes to changes in precipitation showed that none of the ecosystems would shift towards a new type under the range of precipitation increases suggested by the IPCC (increases from 5 to 20%). However, deciduous and semi‐deciduous forests may be very sensitive to small reductions in both the amount and seasonality of precipitation. Main conclusions This version of LPJ‐GUESS parameterized for Africa simulated correctly the vegetation present over a wide precipitation gradient. The biome sensitivity assessment showed that, compared with savannas and grasslands, closed canopy forests may be more sensitive to change in precipitation regime due to the synergetic effects of changed rainfall amounts and seasonality on vegetation functioning.

Abstract The Late Devonian was a period of drastic environmental changes, as exemplified by a major biotic crisis at the Frasnian–Famennian boundary (FFB) and the onset in Famennian times of glaciations across southern Gondwana. Worldwide evidence for the coeval development of the major Acadian–Eovariscan belt led us to propose a model relating the Late Frasnian–Famennian environmental perturbations to extensive continental uplift through two atmospheric CO 2 ‐depleting mechanisms: (1) the intensification of silicate weathering on the continental areas as attested by a major rise in the 87 Sr/ 86 Sr composition of sea water at the FFB; and (2) the massive burial of organic carbon (Kellwasser events) in partially confined basins due to the collisional‐induced reduction of equatorial oceanic communications between the Palaeotethysian and Panthalassic oceans. This process is also suggested to have been controlled by an important primary productivity connected to an increased nutrient availability triggered by the enhanced continental run‐off.

Under a single ‘fault-type hypothesis’, either actuator/component faults or sensor faults, geometric conditions are given such that the original nonlinear system can be transformed into two different subsystems with uncertainty. The first is in the generalised observer canonical form, which is not affected by faults. The second, whose states can be measured, is affected by the faults. Constructing a nonlinear observer for the first subsystem allows estimation of the faults from the second subsystem. Robust (sliding mode) observers are proposed depending on different assumptions on the system uncertainties. The decision logic associated with the proposed bank of observers, and some extensions to simultaneous actuator and sensor faults case are also discussed. Two numerical examples are given to illustrate the design procedures and the efficiency of the proposed methods.

Abstract Aim Poleward range shifts of species are among the most obvious effects of climate change on biodiversity. As a consequence of these range shifts, species communities are predicted to become increasingly composed of warm‐dwelling species, but this has only been studied for a limited number of taxa, mainly birds, butterflies and plants. As species groups may vary considerably in their adaptation to climate change, it is desirable to expand these studies to other groups, from different ecosystems. Freshwater macroinvertebrates, such as dragonflies (Odonata), have been ranked among the species groups with highest priority. In this paper, we investigate how the occurrence of dragonflies in Europe has changed in recent decades, and if these changes are in parallel with climate change. Location Europe. Methods We use data from 10 European geographical regions to calculate occupancy indices and trends for 99 (69%) of the European species. Next, we combine these regional indices to calculate European indices. To determine if changes in regional dragonfly communities in Europe reflect climatic warming, we calculate Species Temperature Indices (STI), Multi‐species Indicators (MSI) and Community Temperature Indices (CTI). Results 55 of 99 considered species increased in occupancy at European level, 32 species remained stable, and none declined. Trends for 12 species are uncertain. MSI of cold‐dwelling and warm‐dwelling species differ in some of the regions, but increased at a similar rate at European level. CTI increased in all regions, except Cyprus. The European CTI increased slightly. Main conclusions European dragonflies, in general, have expanded their distribution in response to climate change, even though their CTI lags behind the increase in temperature. Furthermore, dragonflies proved to be a suitable species group for monitoring changes in communities, both at regional and continental level.

A cDNA sequence with homologies to members of the LPS-binding protein and bactericidal/permeability-increasing protein (BPI) family was identified in the oyster Crassostrea gigas. The recombinant protein was found to bind LPS, to display bactericidal activity against Escherichia coli, and to increase the permeability of the bacterial cytoplasmic membrane. This indicated that it is a BPI rather than an LPS-binding protein. By in situ hybridization, the expression of the C. gigas BPI (Cg-bpi) was found to be induced in hemocytes after oyster bacterial challenge and to be constitutive in various epithelia of unchallenged oysters. Thus, Cg-bpi transcripts were detected in the epithelial cells of tissues/organs in contact with the external environment (mantle, gills, digestive tract, digestive gland diverticula, and gonad follicles). Therefore, Cg-BPI, whose expression profile and biological properties are reminiscent of mammalian BPIs, may provide a first line of defense against potential bacterial invasion. To our knowledge, this is the first characterization of a BPI in an invertebrate.

Abstract. The high-resolution multiproxy study of the Adriatic marine core MD 90-917 provides new insights to reconstruct vegetation and regional climate changes over the southcentral Mediterranean during the Younger Dryas (YD) and Holocene. Pollen records show the rapid forest colonization of the Italian and Balkan borderlands and the gradual installation of the Mediterranean association during the Holocene. Quantitative estimates based on pollen data provide Holocene precipitations and temperatures in the Adriatic Sea using a multi-method approach. Clay mineral ratios from the same core reflect the relative contributions of riverine (illite and smectite) and eolian (kaolinite) contributions to the site, and thus act as an additional proxy with which to evaluate precipitation changes in the Holocene. Vegetation climate reconstructions show the response to the Preboreal oscillation (PBO), most likely driven by changes in temperature and seasonal precipitation, which is linked to increasing river inputs from Adriatic rivers recorded by increase in clay mineral contribution to marine sediments. Pollen-inferred temperature declines during the early–mid Holocene, then increases during the mid–late Holocene, similar to southwestern Mediterranean climatic patterns during the Holocene. Several short vegetation and climatic events appear in the record, indicating the sensitivity of vegetation in the region to millennial-scale variability. Reconstructed summer precipitation shows a regional maximum (170–200 mm) between 8000 and 7000 similar to the general pattern across southern Europe. Two important shifts in vegetation occur at 7700 cal yr BP (calendar years before present) and between 7500 and 7000 cal yr BP and are correlated with increased river inputs around the Adriatic Basin respectively from the northern (7700 event) and from the central Adriatic borderlands (7500–7000 event). During the mid-Holocene, the wet summers lead to permanent moisture all year resulting in a homogeneous seasonal precipitation regime. After 6000 cal yr BP, summer precipitation decreases towards present-day values while winter precipitation rises regularly showing the setting up of Mediterranean climate conditions. Multiproxy evidence from core MD 90-917 provides a deeper understanding of the role of precipitation and particularly the seasonality of precipitation in mediating vegetation change in the central Mediterranean during the Holocene.

Abstract The Ordovician glaciation represents the acme of one of only three major icehouse periods in Earth's Phanerozoic history and is notorious for setting the scene for one of the “big five” mass extinction events. Nevertheless, the mechanisms that drove ice sheet growth remain poorly understood and the final extent of the ice sheet crudely constrained. Here using an Earth system model with an innovative coupling method between ocean, atmosphere, and land ice accounting for climate and ice sheet feedback processes, we report simulations portraying for the first time the detailed evolution of the Ordovician ice sheet. We show that the emergence of the ice sheet happened in two discrete phases. In a counterintuitive sequence of events, the continental ice sheet appeared suddenly in a warm climate. Only during the second act, and set against a background of decreasing atmospheric CO 2 , followed steeply dropping temperatures and extending sea ice. The comparison with abundant sedimentological, geochemical, and micropaleontological data suggests that glacial onset may have occurred as early as the Middle Ordovician Darriwilian, in agreement with recent studies reporting third‐order glacioeustatic cycles during the same period. The second step in ice sheet growth, typified by a sudden drop in tropical sea surface temperatures by ∼8°C and the further extension of a single, continental‐scale ice sheet over Gondwana, marked the onset of the Hirnantian glacial maximum. By suggesting the presence of an ice sheet over Gondwana throughout most of the Middle and Late Ordovician, our models embrace the emerging paradigm of an “early Paleozoic Ice Age.”

K(+) channels in the basolateral membrane of mouse cortical collecting duct (CCD) principal cells were identified with patch-clamp technique, real-time PCR, and immunohistochemistry. In cell-attached membrane patches, three K(+) channels with conductances of approximately 75, 40, and 20 pS were observed, but the K(+) channel with the intermediate conductance (40 pS) predominated. In inside-out membrane patches exposed to an Mg(2+)-free medium, the current-voltage relationship of the intermediate-conductance channel was linear with a conductance of 38 pS. Addition of 1.3 mM internal Mg(2+) had no influence on the inward conductance (G(in) = 35 pS) but reduced outward conductance (G(out)) to 13 pS, yielding a G(in)/G(out) of 3.2. The polycation spermine (6 x 10(-7) M) reduced its activity on inside-out membrane patches by 50% at a clamp potential of 60 mV. Channel activity was also dependent on intracellular pH (pH(i)): a sigmoid relationship between pH(i) and channel normalized current (NP(o)) was observed with a pK of 7.24 and a Hill coefficient of 1.7. By real-time PCR on CCD extracts, inwardly rectifying K(+) (Kir)4.1 and Kir5.1, but not Kir4.2, mRNAs were detected. Kir4.1 and Kir5.1 proteins cellularly colocalized with aquaporin 2 (AQP2), a specific marker of CCD principal cells, while AQP2-negative cells (i.e., intercalated cells) showed no staining. Dietary K(+) had no influence on the properties of the intermediate-conductance channel, but a Na(+)-depleted diet increased its open probability by approximately 25%. We conclude that the Kir4.1/Kir5.1 channel is a major component of the K(+) conductance in the basolateral membrane of mouse CCD principal cells.