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Egbert Giles LEIGH, Jr THE BIOGEOGRAPHY OF LARGE ISLANDS, OR HOW DOES THE SIZE OF THE ECOLOGICAL THEATER AFFECT THE EVOLUTIONARY PLAY? 1 2 2 3 Egbert Giles LEIGH, Jr. , Annette HLADIK , Claude Marcel HLADIK & Alison JOLLY RESUME. — La biogeographie des grandes lies, ou comment la taille de la scene ecologique infiuence- t-elle lejeu de revolution ? —Nous presentons une approche comparative des particularites de revolution dans des milieux insulaires de differentes surfaces, allant de la taille de l'lle de La Reunion a celle de l'Ame- rique du Sud au Pliocene. Cette revue des formes actuelles et fossiles est centree sur Madagascar, ainsi que sur la Nouvelle-Zelande, la Nouvelle-Caledonie et les lies Hawaii, dont les caracteristiques geologiques et historiques sont precisees. L'etendue des terres isolees apparait comme un facteur essentiel qui determine la biodiversite, la taille des plus grands herbivores et celle des predateurs au sommet des chaines trophiques, le rythme de vie (fonc- tion de la longevity et du metabolisme de base des animaux a taille egale), l'intensite de la competition ainsi que la resilience de l'ecosysteme par rapport aux especes envahissantes. Toutes ces caracteristiques dependent aussi de l'eloignement des lies par rapport aux continents. Sur les plus grandes lies isolees pendant de longues periodes, des radiations adaptatives - a partir d'especes colonisatrices ayant pu occasionnellement parcourir une longue distance - peuvent jouer le role ecologique de groupes localement absents, meme si l'anciennete d'une colonisation et la relative protection vis-a-vis des competiteurs continentaux qu'offre le nouvel habitat insulaire ne garantissent pas necessairement une grande diversification des especes qui en derivent. Diversification et endemisme : Bien que Madagascar et la Nouvelle-Zelande aient etc reliees a des blocs continentaux, il y a un peu moins de 90 millions d'annees, leurs biocenoses sont largement dominees par les descendants de formes colonisatrices ayant plus recemment traverse la mer. La phylogenie des especes et leur datation en fonction de l'ADN montrent notamment qu'a Madagascar les formes actuelles de mam- miferes et d'oiseaux, ainsi que presque tous les reptiles et les plantes a fleurs (Tableau I) proviennent des colonisateurs qui ont traverse de vastes etendues marines. Des especes endemiques qui sembleraient ancien- nes comme les baobabs sont arrivees a Madagascar il y a seulement dix millions d'annees ; et la phylogenie moleculaire montre que certains groupes de vertebres terrestres actuellement diversifies en de nombreux genres et especes endemiques (lemuriens, tenrecs), appartiennent chacun, en fait, a une seule radiation, done a une seule colonisation reussie par la forme ancestrale. Inversement, les vagues successives d'especes colo- nisatrices sont illustrees notamment par le hetre austral (Nothofagus) ayant colonise la Nouvelle-Zelande depuis 1'Australie ; alors qu'il avait disparu a la suite de changements climatiques, il a de nouveau colonise ce milieu insulaire. D'une facon generate, les datations en fonction de la phylogenie moleculaire montrent qu'on a longtemps sous-estime l'importance d'especes colonisatrices sur les terres qui furent anciennement separees du continent. Ainsi que l'a montre Darwin, les especes colonisant les terres isolees ont tendance a evoluer vers de nouvelles especes endemiques. La proportion de ces endemiques est plus grande sur les terres les plus iso- lees, par exemple aux iles Hawaii, notre site le plus eloigne, ou le taux d'endemisme est particulierement eleve. Le taux d'endemisme est aussi plus grand dans les groupes dont la dissemination est peu efficace : a Madagascar, ce taux est plus eleve pour les plantes a fleurs que pour les fougeres et egalement plus eleve chez les amphibiens, les reptiles ou les mammiferes terrestres que chez les oiseaux ou les libellules (Tableau II). De plus, un faible nombre de groupes peuvent se differencier sur des iles isolees et occuper des niches eco- logiques qui ne sont pas accessibles aux especes des groupes continentaux apparentes. Les exemples cites a partir des donnees bibliographiques recentes ont permis de preciser les mecanismes de diversification des especes sur les iles et les archipels. La speciation fut allopatrique, aussi bien pour Drosophila sur les iles 1 Smithsonian Tropical Research Institute, Unit 0948, APO AA 34002-0948, USA. Eco-Anthropologie, CNRS and Museum National d'Histoire Naturelle, 4 Avenue du Petit Chateau, F-91800 Brunoy, France. School of Biological Sciences, University of Sussex, Palmer, Brighton BN1 9RH, UK. Rev. Ecol. (Terre Vie), vol. 62, 2007. -105- Hawaii, que pour les arbres de la section Tieghemopanax (Polyscias, Araliaceae) en Nouvelle-Caledonie, ainsi que pour les lemuriens de Madagascar. Cette differenciation des especes correspond generalement aux exigences contrastees des differents habitats et des divers modes de vie possibles. La selection dans les populations en train de se diversities va dans le sens d'une diminution de la competition pour les ressources limitees. Ainsi a Madagascar, les plantules des differentes formes de l'arbre du voyageur (Ravenala) sont respectivement adaptees aux sous-bois, aux espaces degages ou aux milieux marecageux et le decalage des floraisons des formes arborescentes adultes permet la sympatrie dans certains cas. Sur les lies les plus isolees et sur celles d'etendue suffisante pour heberger une grande diversite, on trouve les radiations evolutives les plus spectaculaires telles les drosophiles et les Drepanididae d'Hawaii ou, a Madagascar, les palmiers Dyp- sis, les batraciens Mantellidae et les lemuriens. Selon le modele darwinien, les innovations sont bien plus frequentes sur les terres de grande etendue. Tous les continents sauf l'Antarctique (l'Australie et l'Amerique du Sud independamment des autres) ont permis r emergence des ecosystemes prairiaux (ou gramineens) incluant de grands herbivores. Toutefois a Madagascar, l'hippopotame, un brouteur qui occupa les milieux ouverts et s'y differencia, derive d'une forme venue du continent; et aucune des iles de plus petite surface que Madagascar n'a vu evoluer ce type d'ecosysteme prairial. Diversite, faille, convergence et rythme de vie : La probability d'extinction est plus faible et les possi- bility de diversification sont plus grandes sur les iles de grande surface ; c'est done dans ces milieux que la diversite des especes, locale et/ou totale, est la plus grande (Tableaux VI et VIII). L'analyse des releves bota- niques, par le calcul de l'indice a de Fisher selon la formule S = a ln(l + N/a) ou S est le nombre d'especes et N le nombre des arbres, a permis de mettre en evidence ces differences. Alors qu'en Nouvelle-Caledonie (riche de 3 061 especes indigenes), un echantillon de 500 arbres de foret dense correspond a 76 especes, a Madagascar (11 000 especes indigenes) il correspond a 123 especes et en Nouvelle-Guinee (15 000 especes indigenes) a 196 especes. De la meme facon, les oiseaux insulaires utilisent davantage d'habitats, avec un regime alimentaire plus eclectique que celui de leurs equivalents continentaux. II en resulte une plus grande competition sur les iles de grande surface, alors que la pression des herbivores et des predateurs se fait d'autant moins sentir que les iles sont petites, avec egalement, dans ces cas, une tendance vers une plus faible production primaire et un rythme de vie plus lent. Ainsi, avant l'apparition de l'homme, les plus gros herbivores etaient de moindre taille dans les milieux insulaires et d'autant plus petits que la surface des iles etait reduite. Alors que les elephants et les mam- mouths depassaient 5 tonnes sur les blocs continentaux de grande surface, l'oiseau-elephant de Madagascar (Aepyornis) pesait 275 kg et le plus gros herbivore d'Hawaii seulement 8,6 kg. La productivite primaire des ecosystemes est actuellement plus faible la ou ces consommateurs primaires etaient de plus petite taille, notamment a Hawaii et probablement aussi a Madagascar (Tableau XIV). Parallelement il apparait qu'il y a moins d'especes pionnieres efficaces pour la regeneration forestiere sur les iles que dans les forets conti- nentales. Cela pourrait s'expliquer par le fait que, sur les iles, les plus gros folivores ne pouvaient pas creer de nombreux chablis en faisant chuter les arbres (comme les elephants peuvent le faire). Les carnivores sont egalement de moindre taille dans les milieux insulaires de surface reduite, compares aux grands felins d'Afrique ou d'Asie, avec seulement 17 kg pour Cryptoprocta spelea a Madagascar et a Hawaii 5 kg pour 1'aigle qui etait localement le plus gros des predateurs (Tableau XIII). Dans la mesure ou, sur les iles, les especes sont exposees, au stade adulte, a de moindres dangers de predation, elles peuvent consacrer une plus grande partie des ressources a un allongement de la duree de vie et moins investir pour se reproduire rapidement et en grand nombre. Ces particularity des milieux insulaires furent observees et decrites chez les rongeurs par Adler et Levins, sous le nom de " syndrome ilien ". Elles s'observent notamment chez le genre Anolis qui, sur les iles Cara'ibes, vit plus longtemps et se reproduit plus tardivement et en moindre nombre que les especes continentales equivalentes, ainsi que chez de nombreuses especes de Nouvelle-Zelande reputees pour leur longevite et leur reproduction extremement lente. La faible pression de predation se traduit egalement, chez les herbivores, par des convergences de formes et de fonc- tions entre des especes insulaires et continentales la ou les predateurs sont peu efficaces. Par exemple, des lemuriens fossiles presenters des convergences avec des paresseux terrestres ou arboricoles d'Amerique du Sud et avec les koalas d'Australie, qui vivaient dans un environnement ou les predateurs marsupiaux etaient relativement peu efficaces. De meme, les moas (Dinornis) de Nouvelle-Zelande ont evolue de facon conver- gente avec les moa-nalos (Branta et Chelychelynechen) des iles Hawaii et les oiseaux-elephants (Aepyornis) de Madagascar.
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