Alain Dejean

French National Centre for Scientific Research, Lutetia Parisorum, Île-de-France, France

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Publications (234)445.13 Total impact

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    ABSTRACT: RationaleCompared with other animal venoms, ant venoms remain little explored. Ants have evolved complex venoms to rapidly immobilize arthropod prey and to protect their colonies from predators and pathogens. Many ants have retained peptide-rich venoms that are similar to those of other arthropod groups.Methods With the goal of conducting a broad and comprehensive survey of ant venom peptide diversity, we investigated the peptide composition of venoms from 82 stinging ant species from nine subfamilies using matrix-assisted laser desorption/ionisation time-of-flight mass spectrometry (MALDI-TOFMS). We also conducted an in-depth investigation of eight venoms using reversed-phase high-performance liquid chromatography (RP-HPLC) separation coupled with offline MALDI-TOFMS.ResultsOur results reveal that the peptide compositions of ant venom peptidomes from both poneroid and formicoid ant clades comprise hundreds of small peptides (<4 kDa), while large peptides (>4 kDa) are also present in the venom of formicoids. Chemical reduction revealed the presence of disulfide-linked peptides in most ant subfamilies, including peptides structured by one, two or three disulfide bonds as well as dimeric peptides reticulated by three disulfide bonds.Conclusions The biochemical complexity of ant venoms, associated with an enormous ecological and taxonomic diversity, suggests that stinging ant venoms constitute a promising source of bioactive molecules that could be exploited in the search for novel drug and biopesticide leads. Copyright © 2015 John Wiley & Sons, Ltd.
    Rapid Communications in Mass Spectrometry 03/2015; 29(5). DOI:10.1002/rcm.7116 · 2.51 Impact Factor
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    ABSTRACT: We show that in French Guiana the large carton nests of Azteca chartifex, a territorially-dominant arboreal dolichoderine ant, are protected from bird attacks when this ant lives in association with Polybia rejecta, an epiponine social wasp. Because A. chartifex colonies are well known for their ability to divert army ant raids from the base of their host tree so that they protect their associated wasps from these raids, there is a reciprocal benefit for these two partners, permitting us to call this association a mutualism. We also show that P. rejecta nests are significantly less often attacked by birds than are those of two compared epiponine social wasp species. Furthermore, experimentation using a standardized protocol demonstrated the significantly higher aggressiveness of P. rejecta compared to seven other wasp species. We conclude that the efficacious protection of its associated ant nests is likely due to the extreme aggressiveness of P. rejecta.
    Comptes Rendus Biologies 03/2015; Online first. · 1.68 Impact Factor
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    ABSTRACT: We show that in French Guiana the large carton nests of Azteca chartifex, a territorially-dominant arboreal dolichoderine ant, are protected from bird attacks when this ant lives in association with Polybia rejecta, an epiponine social wasp. Because A. chartifex colonies are well known for their ability to divert army ant raids from the base of their host tree so that they protect their associated wasps from these raids, there is a reciprocal benefit for these two partners, permitting us to call this association a mutualism. We also show that P. rejecta nests are significantly less often attacked by birds than are those of two compared epiponine social wasp species. Furthermore, experimentation using a standardized protocol demonstrated the significantly higher aggressiveness of P. rejecta compared to seven other wasp species. We conclude that the efficacious protection of its associated ant nests is likely due to the extreme aggressiveness of P. rejecta. Copyright © 2015 Académie des sciences. Published by Elsevier SAS. All rights reserved.
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    ABSTRACT: Ants, the most abundant taxa among canopy-dwelling animals in tropical rainforests, are mostly represented by territorially-dominant arboreal ants (TDAs) whose territories are distributed in a mosaic pattern (arboreal ant mosaics). Large TDA colonies regulate insect herbivores, with implications for forestry and agronomy. What generates these mosaics in vegetal formations, which are dynamic, still needs to be better understood. So, from empirical research based on three Cameroonian tree species (Lophira alata, Ochnaceae; Anthocleista vogelii, Gentianaceae; and Barteria fistulosa, Passifloraceae), we used the Self-Organizing Map (SOM, neural network) to illustrate the succession of TDAs as their host trees grow and age. The SOM separated the trees by species and by size for L. alata, which can reach 60 m in height and live several centuries. An ontogenic succession of TDAs from sapling to mature trees is shown, and some ecological traits are highlighted for certain TDAs. Also, because the SOM permits the analysis of data with many zeroes with no effect of outliers on the overall scatterplot distributions, we obtained ecological information on rare species. Finally, the SOM permitted us to show that functional groups cannot be selected at the genus level as congeneric species can have very different ecological niches, something particularly true for Crematogaster spp. which include a species specifically associated with B. fistulosa, non-dominant species and TDAs. Therefore, the SOM permitted the complex relationships between TDAs and their growing host trees to be analyzed, while also providing new information on the ecological traits of the ant species involved. This article is protected by copyright. All rights reserved. This article is protected by copyright. All rights reserved.
    Insect Science 02/2015; DOI:10.1111/1744-7917.12208 · 1.51 Impact Factor
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    ABSTRACT: Tank bromeliads are good models for understanding how climate change may affect biotic associations. We studied the relationships between spiders, the epiphytic tank bromeliad, Aechmea bracteata, and its associated ants in an inundated forest in Quintana Roo, Mexico, during a drought period while, exceptionally, this forest was dry and then during the flooding that followed. We compared spider abundance and diversity between ‘Aechmea-areas’ and ‘control-areas’ of the same surface area. We recorded six spider families: the Dipluridae, Ctenidae, Salticidae, Araneidae, Tetragnathidae and Linyphiidae among which the funnel-web tarantula, Ischnothele caudata, the only Dipluridae noted, was the most abundant. During the drought period, the spiders were more numerous in the Aechmea-areas than in the control-areas, but they were not obligatorily associated with the Aechmea. During the subsequent flooding, the spiders were concentrated in the A. bracteata patches, particularly those sheltering an ant colony. Also, a kind of specificity existed between certain spider taxa and ant species, but varied between the drought period and subsequent flooding. We conclude that climatic events modulate the relationship between A. bracteata patches and their associated fauna. Tank bromeliads, previously considered only for their ecological importance in supplying food and water during drought, may also be considered refuges for spiders during flooding. More generally, tank bromeliads have an important role in preserving non-specialized fauna in inundated forests.
    PLoS ONE 12/2014; DOI:10.1371/journal.pone.0114592 · 3.53 Impact Factor
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    Alain Dejean, Bruno Corbara
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    ABSTRACT: Il a été montré dernièrement que les colonies de la fourmi invasive Pheidole megacephala contrattaquent lors de raids par les fourmis légionnaires Eciton burchellii. Les contacts qui en résultent permettent le transfert de composés cuticulaires (ils constituent l’odeur coloniale) des Pheidole vers les Eciton. Ces dernières ne sont plus reconnues par leurs congénères, qui les tuent. Utilisant une méthode simple pour transférer des composés cuticulaires, nous avons testé si ce phénomène existait pour des fourmis néotropicales. Des Eciton frottées avec des ouvrières Solenopsis saevissima ou Camponotus blandus ont été attaquées par leurs congénères, mais pas celles frottées avec des Atta sexdens, des Pheidole fallax ou des congénères (lot contrôle). Ainsi, les composés cuticulaires de certaines espèces sympatriques déclenchent l’agressivité entre ouvrières Eciton s’ils sont transférés, alors que d’autres sont sans effet. Nous concluons que ces composés pourraient avoir joué un rôle dans l’évolution du comportement prédateur chez les fourmis légionnaires.
    Comptes Rendus Biologies 11/2014; DOI:10.1016/j.crvi.2014.08.008 · 1.68 Impact Factor
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    ABSTRACT: Ants (Hymenoptera: Formicidae) represent a taxonomically diverse group of arthropods comprising nearly 13,000 extant species. Sixteen ant subfamilies have individuals that possess a stinger and use their venom for purposes such as a defence against predators, competitors and microbial pathogens, for predation, as well as for social communication. They exhibit a range of activities including antimicrobial, haemolytic, cytolytic, paralytic, insecticidal and pain-producing pharmacologies. While ant venoms are known to be rich in alkaloids and hydrocarbons, ant venoms rich in peptides are becoming more common, yet remain understudied. Recent advances in mass spectrometry techniques have begun to reveal the true complexity of ant venom peptide composition. In the few venoms explored thus far, most peptide toxins appear to occur as small polycationic linear toxins, with antibacterial properties and insecticidal activity. Unlike other venomous animals, a number of ant venoms also contain a range of homodimeric and heterodimeric peptides with one or two interchain disulfide bonds possessing pore-forming, allergenic and paralytic actions. However, ant venoms seem to have only a small number of monomeric disulfide-linked peptides. The present review details the structure and pharmacology of known ant venom peptide toxins and their potential as a source of novel bioinsecticides and therapeutic agents.
    Toxicon 10/2014; DOI:10.1016/j.toxicon.2014.10.021 · 2.58 Impact Factor
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    ABSTRACT: The phenotypic plasticity of plants has been explored as a function of either ontogeny (apparent plasticity) or environment (adaptive plasticity), although few studies have analyzed these factors together. In the present study, we take advantage of the dispersal of Aechmea mertensii bromeliads by Camponotus femoratus or Pachycondyla goeldii ants in shaded and sunny environments, respectively, to quantify ontogenetic changes in morphological, foliar, and functional traits, and to analyze ontogenetic and ant species effects on 14 traits. Most of the morphological (plant height, number of leaves), foliar (leaf thickness, leaf mass area, total water content, trichome density), and functional (leaf δ13C) traits differed as a function of ontogeny. Conversely, only leaf δ15N showed an adaptive phenotypic plasticity. On the other hand, plant width, tank width, longest leaf length, stomatal density, and leaf C concentration showed an adaptation to local environment with ontogeny. The exception was leaf N concentration, which showed no trend at all. Aechmea mertensii did not show an abrupt morphological modification such as in heteroblastic bromeliads, although it was characterized by strong, size-related functional modifications for CO2 acquisition. The adaptive phenotypic variation found between the two ant species indicates the spatially conditioned plasticity of A. mertensii in the context of insect-assisted dispersal. However, ant-mediated effects on phenotypic plasticity in A. mertensii are not obvious because ant species and light environment are confounding variables. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, ●●, ●●–●●.
    Botanical Journal of the Linnean Society 06/2014; 175(2). DOI:10.1111/boj.12171 · 2.70 Impact Factor
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    ABSTRACT: We tested if nesting habits influence ant feeding preferences and predatory behavior in the monophyletic genus Pseudomyrmex (Pseudomyrmecinae) which comprises terrestrial and arboreal species, and, among the latter, plant-ants which are obligate inhabitants of myrmecophytes (i.e., plants sheltering so-called plant-ants in hollow structures). A cafeteria experiment revealed that the diet of ground-nesting Pseudomyrmex consists mostly of prey and that of arboreal species consists mostly of sugary substances, whereas the plant-ants discarded all the food we provided. Workers forage solitarily, detecting prey from a distance thanks to their hypertrophied eyes. Approach is followed by antennal contact, seizure, and the manipulation of the prey to sting it under its thorax (next to the ventral nerve cord). Arboreal species were not more efficient at capturing prey than were ground-nesting species. A large worker size favors prey capture. Workers from ground- and arboreal-nesting species show several uncommon behavioral traits, each known in different ant genera from different subfamilies: leaping abilities, the use of surface tension strengths to transport liquids, short-range recruitment followed by conflicts between nestmates, the consumption of the prey's hemolymph, and the retrieval of entire prey or pieces of prey after having cut it up. Yet, we never noted group ambushing. We also confirmed that Pseudomyrmex plant-ants live in a kind of food autarky as they feed only on rewards produced by their host myrmecophyte, or on honeydew produced by the hemipterans they attend and possibly on the fungi they cultivate.
    Naturwissenschaften 02/2014; 101(4). DOI:10.1007/s00114-014-1159-1 · 1.97 Impact Factor
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    ABSTRACT: The rise of integrative taxonomy, a multi-criteria approach used in characterizing species, fosters the development of new tools facilitating species delimitation. Mass spectrometric (MS) analysis of venom peptides from venomous animals has previously been demonstrated to be a valid method for identifying species. Here we aimed to develop a rapid chemotaxonomic tool for identifying ants based on venom peptide mass fingerprinting. The study focused on the biodiversity of ponerine ants (Hymenoptera: Formicidae: Ponerinae) in French Guiana. Initial experiments optimized the use of automated matrix-assisted laser desorption/ionization time-of-flight mass spectrometry (MALDI-TOF MS) to determine variations in the mass profiles of ant venoms using several MALDI matrices and additives. Data were then analyzed via a hierarchical cluster analysis to classify the venoms of 17 ant species. In addition, phylogenetic relationships were assessed and were highly correlated with methods using DNA sequencing of the mitochondrial gene cytochrome c oxidase subunit 1. By combining a molecular genetics approach with this chemotaxonomic approach, we were able to improve the accuracy of the taxonomic findings to reveal cryptic ant species within species complexes. This chemotaxonomic tool can therefore contribute to more rapid species identification and more accurate taxonomies. This is the first extensive study concerning the peptide analysis of the venom of both Pachycondyla and Odontomachus ants. We studied the venoms of 17 ant species from French Guiana that permitted us to fine-tune the venom analysis of ponerine ants via MALDI-TOF mass spectrometry. We explored the peptidomes of crude ant venom and demonstrated that venom peptides can be used in the identification of ant species. In addition, the application of this novel chemotaxonomic method combined with a parallel genetic approach using COI sequencing permitted us to reveal the presence of cryptic ants within both the Pachycondyla apicalis and Pachycondyla stigma species complexes. This adds a new dimension to the search for means of exploiting the enormous biodiversity of venomous ants as a source for novel therapeutic drugs or biopesticides.
    Journal of proteomics 01/2014; DOI:10.1016/j.jprot.2014.01.009 · 5.07 Impact Factor
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    ABSTRACT: The tank bromeliads Aechmea aquilega (Salisb.) and Catopsis berteroniana (Schultes f.) coexist on a sun-exposed Neotropical inselberg in French Guiana, where they permit conspicuous freshwater pools to form that differ in size, complexity and detritus content. We sampled the algal communities (both eukaryotic and cyanobacterial taxa, including colourless forms) inhabiting either A. aquilega (n = 31) or C. berteroniana (n = 30) and examined differences in community composition and biomass patterns in relation to several biotic and abiotic variables. Chlorella sp. and Bumilleriopsis sp. were the most common taxa and dominated the algal biomass in A. aquilega and C. berteroniana, respectively. Using a redundancy analysis, we found that water volume, habitat complexity and the density of phagotrophic protozoa and collector-gatherer invertebrates were the main factors explaining the distribution of the algal taxa among the samples. Hierarchical clustering procedures based on abundance and presence/absence data clearly segregated the samples according to bromeliad species, revealing that the algal communities in the smaller bromeliad species were not a subset of the communities found in the larger bromeliad species. We conclude that, even though two coexisting tank bromeliad populations create adjacent aquatic habitats, each population hosts a distinct algal community. Hence, bromeliad diversity is thought to promote the local diversity of freshwater algae in the Neotropics.
    Plant Biology 01/2014; 16(5). DOI:10.1111/plb.12139 · 2.32 Impact Factor
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    ABSTRACT: Supercolonies of the red fire ant Solenopsis saevissima (Smith) develop in disturbed environments and likely alter the ant community in the native range of the species. For example, in French Guiana only eight ant species were repeatedly noted as nesting in close vicinity to its mounds. Here, we verified if a shared set of biological, ecological and behavioral traits might explain how these eight species are able to nest in the presence of S. saevissima. We did not find this to be the case. We did find, however, that all of them are able to live in disturbed habitats. It is likely that over the course of evolution each of these species acquired the capacity to live syntopically with S. saevissima through its own set of traits, where colony size (four species develop large colonies), cuticular compounds which do not trigger aggressiveness (six species) and submissive behaviors (four species) complement each other.
    Myrmecological News 01/2014; 19:17-24. · 2.16 Impact Factor
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    Myrmecological News 01/2014; 19:43-51. · 2.16 Impact Factor
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    ABSTRACT: Zelus annulosus is an assassin bug species mostly 28 noted on Hirtella physophora, a myrmecophyte specifically associated with the ant Allomerus decemarticulatus known to build traps on host tree twigs to ambush insect prey. The females lay egg clutches protected by a sticky substance. To avoid being trapped, the first three instars of Z. annulosus nymphs remain grouped in a clutch beneath the leaves where they hatched, yet from time to time they climb onto the upper side to group ambush prey. Long-distance prey detection permits these bugs to capture flying or jumping insects that alight on their leaves. Like for some other Zelus species, the sticky substance of the sundew setae of their forelegs aids in prey capture. Group ambushing permits early instars to capture insects that they then share or not depending on prey size and the hunger of the successful nymphs. Fourth and fifth instars, with greater needs, rather ambush solitarily on different host tree leaves, but attract siblings to share large prey. Communal feeding permits faster prey consumption, enabling small nymphs to return sooner to the shelter of their leaves. By improving the regularity of feeding for each nymph, it likely regulates nymphal development, synchronizing molting and subsequently limiting cannibalism.
    Naturwissenschaften 10/2013; 100:913-922. DOI:10.1007/s00114-013-1091-9 · 1.97 Impact Factor
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    ABSTRACT: Melissotarsus ants have an extremely specialized set of behaviours. Both workers and gynes tunnel galleries in their host tree bark. Workers walk with their mesothoracic legs pointing upwards and tend Diaspididae hemiptera for their flesh. The ants use their forelegs to plug the galleries with silk that they secrete themselves. We hypothesised that the ants' energetic needs for nearly constant gallery digging could be satisfied through the absorption of host tree tissues; so, using basic techniques, we examined the digestive capacities of workers from two species. We show that workers are able to degrade oligosaccharides and heterosides as well as, to a lesser degree, polysaccharides. This is one of the rare reports on ants able to digest plant polysaccharides other than starch.
    Comptes rendus biologies 10/2013; 336(10):500-4. DOI:10.1016/j.crvi.2013.08.003 · 1.71 Impact Factor
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    ABSTRACT: Little is known of how linkage patterns between species change along environmental gradients. The small, spatially discrete food webs inhabiting tank-bromeliads provide an excellent opportunity to analyse patterns of community diversity and food-web topology (connectance, linkage density, nestedness) in relation to key environmental variables (habitat size, detrital resource, incident radiation) and predators:prey ratios. We sampled 365 bromeliads in a wide range of understorey environments in French Guiana and used gut contents of invertebrates to draw the corresponding 365 connectance webs. At the bromeliad scale, habitat size (water volume) determined the number of species that constitute food-web nodes, the proportion of predators, and food-web topology. The number of species as well as the proportion of predators within bromeliads declined from open to forested habitats, where the volume of water collected by bromeliads was generally lower because of rainfall interception by the canopy. A core group of microorganisms and generalist detritivores remained relatively constant across environments. This suggests that (i) a highly-connected core ensures food-web stability and key ecosystem functions across environments, and (ii) larger deviations in food-web structures can be expected following disturbance if detritivores share traits that determine responses to environmental changes. While linkage density and nestedness were lower in bromeliads in the forest than in open areas, experiments are needed to confirm a trend for lower food-web stability in the understorey of primary forests.
    PLoS ONE 08/2013; 8(8):e71735. DOI:10.1371/journal.pone.0071735 · 3.53 Impact Factor
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    ABSTRACT: Background and AimsEpiphytism imposes physiological constraints resulting from the lack of access to the nutrient sources available to ground-rooted plants. A conspicuous adaptation in response to that lack is the phytotelm (plant-held waters) of tank-bromeliad species that are often nutrient-rich. Associations with terrestrial invertebrates also result in higher plant nutrient acquisition. Assuming that tank-bromeliads rely on reservoir-assisted nutrition, it was hypothesized that the dual association with mutualistic ants and the phytotelm food web provides greater nutritional benefits to the plant compared with those bromeliads involved in only one of these two associations.Methods Quantitative (water volume, amount of fine particulate organic matter, predator/prey ratio, algal density) and qualitative variables (ant-association and photosynthetic pathways) were compared for eight tank- and one tankless-bromeliad morphospecies from French Guiana. An analysis was also made of which of these variables affect nitrogen acquisition (leaf N and δ(15)N).Key ResultsAll variables were significantly different between tank-bromeliad species. Leaf N concentrations and leaf δ(15)N were both positively correlated with the presence of mutualistic ants. The amount of fine particulate organic matter and predator/prey ratio had a positive and negative effect on leaf δ(15)N, respectively. Water volume was positively correlated with leaf N concentration whereas algal density was negatively correlated. Finally, the photosynthetic pathway (C3 vs. CAM) was positively correlated with leaf N concentration with a slightly higher N concentration for C3-Tillandsioideae compared with CAM-Bromelioideae.Conclusions The study suggests that some of the differences in N nutrition between bromeliad species can be explained by the presence of mutualistic ants. From a nutritional standpoint, it is more advantageous for a bromeliad to use myrmecotrophy via its roots than to use carnivory via its tank. The results highlight a gap in our knowledge of the reciprocal interactions between bromeliads and the various trophic levels (from bacteria to large metazoan predators) that intervene in reservoir-assisted nutrition.
    Annals of Botany 07/2013; DOI:10.1093/aob/mct147 · 3.45 Impact Factor
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    ABSTRACT: In this study, which was carried out on 460 ant-gardens found in an orchard of Citrus grandis and in two pioneer formations of different ages, we noted that four ant species, Pachycondyla goeldii (Ponerinae), Crematogaster limata parabiotica (Myrmicinae), Dolichoderus bispinosus (Dolichoderinae) and Camponotus femoratus (Formicinae), built carton nests on which we found epiphytes at different stages of development. For other ant species sheltering in ant gardens, we did not find young, developing epiphytes. A three-way relationship between the supporting bush, epiphyte and ant becomes apparent. The diversity of the epiphytes varies as a function of the ant species: Aechmea mertensii (Bromeliaceae) is dominant in the gardens of P. goeldii; Codonanthe crassifolia (Gesneriaceae) and Anthurium gracile (Araceae) in those of C. femoratus and of Cr. limata parabiotica, both of which also associated with Peperomia macrostachya (Piperaceae).
    Acta botanica Gallica: bulletin de la Société botanique de France 04/2013; 144(3):333-345. DOI:10.1080/12538078.1997.10515378 · 0.24 Impact Factor
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    ABSTRACT: This article documents the addition of 268 microsatellite marker loci to the Molecular Ecology Resources Database. Loci were developed for the following species: Alburnoides bipunctatus, Chamaerops humilis, Chlidonias hybrida, Cyperus papyrus, Fusarium graminearum, Loxigilla barbadensis, Macrobrachium rosenbergii, Odontesthes bonariensis, Pelteobagrus vachelli, Posidonia oceanica, Potamotrygon motoro, Rhamdia quelen, Sarotherodon melanotheron heudelotii, Sibiraea angustata, Takifugu rubripes, Tarentola mauritanica, Trimmatostroma sp. and Wallago attu. These loci were cross-tested on the following species: Alburnoides fasciatus, Alburnoides kubanicus, Alburnoides maculatus, Alburnoides ohridanus, Alburnoides prespensis, Alburnoides rossicus, Alburnoides strymonicus, Alburnoides thessalicus, Alburnoides tzanevi, Carassius carassius, Fusarium asiaticum, Leucaspius delineatus, Loxigilla noctis dominica, Pelecus cultratus, Phoenix canariensis, Potamotrygon falkneri, Trachycarpus fortune and Vimba vimba.
    Molecular Ecology Resources 03/2013; 13:546-549. DOI:10.1111/1755-0998.12095 · 7.43 Impact Factor

Publication Stats

2k Citations
445.13 Total Impact Points

Institutions

  • 2001–2015
    • French National Centre for Scientific Research
      • Laboratoire d'Ecologie Fonctionnelle et Environnement
      Lutetia Parisorum, Île-de-France, France
    • Tel Aviv University
      Tell Afif, Tel Aviv, Israel
  • 1998–2014
    • University of Toulouse
      Tolosa de Llenguadoc, Midi-Pyrénées, France
  • 1999–2013
    • Paul Sabatier University - Toulouse III
      • • Laboratoire Evolution et Diversité Biologiques (EDB)
      • • Centre de Recherches sur la Cognition Animale - UMR 5169 - CRCA
      Tolosa de Llenguadoc, Midi-Pyrénées, France
  • 2012
    • Smithsonian Tropical Research Institute
      Ciudad de Panamá, Panamá, Panama
  • 2008
    • Lower Columbia College
      Longview, Texas, United States
  • 2006–2008
    • Le laboratoire évolution et diversité biologique
      Tolosa de Llenguadoc, Midi-Pyrénées, France
  • 2005
    • University of Illinois, Urbana-Champaign
      • Department of Animal Sciences
      Urbana, Illinois, United States
  • 2004
    • Université Blaise Pascal - Clermont-Ferrand II
      • Laboratoire Microorganismes : Génome et Environnement
      Clermont-Ferrand, Auvergne, France
  • 2003
    • University of Douala
      • Department of Biology of Animal Organisms
      Douala, Littoral Region, Cameroon
  • 1994–2000
    • Université Paris 13 Nord
      • LEEC Laboratoire d'Ethologie Expirementale et Comparee
      Île-de-France, France
  • 1992
    • University of Quintana Roo
      Cozumel, Quintana Roo, Mexico