CHAPITRE 5 : Conclusion générale, bilan & perspectives
4. Q UELQUES PERSPECTIVES
La caractérisation moléculaire d’un complexe d’espèces ne peut être exhaustive, ne serait-ce que par les difficultés d’échantillonnage. La situation sécuritaire déjà délicate, ne fait que s’accroître au Mexique depuis une dizaine d’années et ne permet pas pour l’heure, de se rendre sans crainte dans les zones frontalières avec les Etats-Unis (Etats du Nord, dans lesquels on retrouve L. montanus ssp. glabrior et L. montanus ssp. montesii) et avec le Guatemala (Etats du Chiapas, dans lequel on retrouve L. kellermanianus et L. montanus var. austrovolcanicus). Néanmoins, l’inclusion de ces taxons du complexe L. montanus associée à un échantillonnage parmi des espèces moins étroitement liées comme les espèces pérennes des Rocheuses (e.g. L.
lepidus et L. albifrons) permettraient de calibrer davantage la variation génétique et
phénotypique à l’intérieur de L. montanus. Sans nul doute, ce plus large échantillonnage permettrait de tester l’hypothèse de monophylie du complexe avec robustesse. Au vu de nos résultats, il semblerait que l’hypothèse monophylétique soit privilégiée, mais l’inclusion de l’ensemble des taxons L. montanus et d’espèces externes plus éloignées dans les analyses apporterait une réponse plus claire.
Malgré un postulat de départ clairement axé sur le manque de résolution des caractères morphologiques dans la détermination des taxons du complexe L. montanus, il reste essentiel de combiner aux patrons moléculaires d’autres types de données parmi lesquelles les préférences écologiques et les caractères morphologiques. En effet, la récente prolifération des études portant sur des complexes d’espèces de plantes ou d’autres groupes d’organismes est généralement basée sur la découverte de différenciation génétique basée sur des données moléculaires et sur des données morphologiques et écologiques. Ainsi, le choix restreint de traits morphologiques « objectifs » apporterait des éléments corroboratifs aux données génétiques
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pour clarifier les relations de parenté à l’intérieur du complexe. Plutôt que des données strictement métriques (peu recommandables dans les études populationnelles ou à faible échelle taxinomique), la conception de matrices morphologiques basées sur des rapports de variables morphométriques semble mieux adaptée, afin d’éviter les problèmes d’analyse et de statistiques liés à l’effet taille. Il faudrait distinguer les vraies différences quantitatives (divergence et singularité morphologique) des fausses différences (notamment basées sur des adaptations locales aux conditions abiotiques générant par exemple des tailles plus modestes en haute altitude).
La collecte de données écologiques, notamment autour des préférences pédologiques et édaphologiques de chaque population et taxon, fournirait des variables supplémentaires qui complèteraient les variables bioclimatiques nécessaires à la modélisation des distributions de niches. Assez peu utilisés, les tests d’identité de niches sont fiables pour savoir s’il y a ou non, chevauchement des préférences écologiques. Etant donné que les modèles actuels de distribution de niches ne font aucune hypothèse biologique concernant l’utilisation de micro- habitats ou l’existence d’interactions entre taxons, des auteurs20 ont récemment proposé une approche mathématique qui note la similarité de niches de 0 (pas de chevauchement de niches) à 1 (chevauchement complet). Ce type de test semble tout à fait adapté à l’échelle populationnelle et permettrait de confirmer l’absence de spéciation écologique chez L. montanus avec beaucoup plus de précision.
Un échantillonnage plus dense requiert de fait, une plus grande connaissance des marqueurs moléculaires utilisés, tant pour éviter un poids trop important des homoplasies que pour éviter des effets d’incongruence liés à la saturation des séquences utilisées. L’ajout d’espèces moins apparentées, pour tester notamment l’hypothèse monophylétique, devra être fait sous condition de vérifier l’état de saturation des marqueurs utilisés. Lorsque deux sites de deux taxons portent le même nucléotide, il n’est pas possible a priori de savoir s’il y a eu 0, 2 ou plus de mutations dans l’histoire de cette séquence. Il n’est pas possible également de déterminer si l’identité de séquence est le fruit d’une convergence (c’est-à-dire une succession de mutations qui provoque un retour à l’état initial, générant un jeu de données qualifié de « saturé ») ou si la séquence demeure inchangée. Le phénomène de saturation, qui peut se calculer en comparant le nombre de différences observées (inférées par les modèles analytiques choisis) et le nombre réel de mutations : plus la pente de la courbe est faible, plus le niveau de saturation est important21.
20 Voir les travaux conduits par Warren et al. (2008).
21 Plus en détail, le taux de saturation apparaît comme étant généralement plus important sur le troisième nucléotide
des codons (une pente moyenne de 0.16), les deux premières positions étant moins sujettes à des substitutions multiples (pente moyenne de 0.43) (Jeffroy et al., 2006).
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