Life‐history traits of Encarsia guadeloupae,a natural enemy of the invasive spiralling whitefly Aleurodicus dispersus

On south‐west Indian Ocean islands, many crops and ornamental plants are threatened by the spiralling whitefly Aleurodicus dispersus (Hemiptera: Aleyrodidae), which is a polyphagous pest that is native to the Caribbean region. Aleurodicus dispersus causes economic damage to various crops on all the islands in the south‐west Indian Ocean. The hymenopteran parasitoid Encarsia guadeloupae (Hymenoptera: Aphelinidae) is a natural enemy of A. dispersus on the Caribbean islands. In this study, we assessed the geographical distribution of the parasitoid in La Réunion, an island in the south‐west Indian Ocean where the parasitoid was first observed in 2004. We also investigated its main life‐history traits. Field surveys indicated that the parasitoid is widespread in most of the low‐lying areas of the island and exhibits high parasitism rates on A. dispersus populations. At 25°C, E. guadeloupae adults had a mean longevity of 33.6 days, and its pre‐imaginal development required 23 days. The lower temperature threshold and thermal constant were estimated to be 7.9°C and 132 degree‐days, respectively. Females of E. guadeloupae preferred to deposit eggs in early rather than in late instars of A. dispersus, and oviposition rates were highest in the second larval instar. Females of E. guadeloupae were able to oviposit in larvae of other species of whiteflies found in La Réunion (Bemisia tabaci and Dialeurolonga simplex), although subsequent development of the parasitoid was not monitored. Finally, we discuss the potential use of E. guadeloupae for the control of whitefly populations on islands in the south‐west Indian Ocean.     

First record of troglobitic Hemiptera (Fulgoromorpha: Cixiidae) from La Réunion Island

A new species of troglobitic Cixiidae (Hemiptera: Fulgoromorpha), Brixia briali n. sp. is described from a lava tube on La Réunion Island. Information on its ecology, distribution, and comments on its tentative systematic placement into the genus Brixia is provided. The new discovery from La Réunion is placed into the context of cavernicolous Fulgoromorpha of the world, their zoogeography, ecology and biology.     

Multi locus analysis of Pristionchus pacificus on La Réunion Island reveals an evolutionary history shaped by multiple introductions, constrained dispersal events and rare out-crossing

Pristionchus pacificus, recently established as a model organism in evolutionary biology, is a cosmopolitan nematode that has a necromenic association with scarab beetles. The diverse array of host beetle species and habitat types occupied by P. pacificus make it a good model for investigating local adaptation to novel environments. Presence of P. pacificus on La Réunion Island, a young volcanic island with a dynamic geological history and a wide variety of ecozones, facilitates such investigation in an island biogeographic setting. Microsatellite data from 20 markers and 223 strains and mitochondrial sequence data from 272 strains reveal rich genetic diversity among La Réunion P. pacificus isolates, shaped by differentially timed introductions from diverse sources and in association with different beetle species. Distinctions between volcanic zones and between arid western and wet eastern climatic zones have likely limited westward dispersal of recently colonized lineages and maintained a genetic distinction between eastern and western clades. The highly selfing lifestyle of P. pacificus contributes to the strong fine‐scale population structure detected, with each beetle host harbouring strongly differentiated assemblages of strains. Periodic out‐crossing generates admixture between genetically diverse lineages, creating a diverse array of allelic combinations likely to increase the evolutionary potential of the species and facilitate adaptation to local environments and beetle hosts.     

Haplotype diversity of the nematode Pristionchus pacificus on Réunion in the Indian Ocean suggests multiple independent invasions

Pristionchus pacificus has been established as a nematode model system in evolutionary developmental biology and evolutionary ecology. Field studies in North and South America, Asia, Africa and Europe indicated that nematodes of the genus Pristionchus live in association with scarab beetles. Here, we describe the first account of soil‐ and beetle‐associated nematodes on an island setting by investigating the island of Réunion in the Indian Ocean. Réunion has high numbers of endemic insects and is one among several attractive islands for biodiversity studies. Being of volcanic origin, Réunion is 2–3 million years old, making it the youngest of the Mascareigne islands. We show that beetle‐ and soil‐derived nematodes on Réunion are nearly exclusively hermaphroditic, suggesting that selfing is favoured over gonochorism (outcrossing) during island colonization. Among members of four nematode genera observed on Réunion, Pristionchus pacificus was the most prevalent species. A total of 76 isolates, in association with five different scarab beetles, has been obtained for this cosmopolitan nematode. A detailed mitochondrial haplotype analysis indicates that the Réunion isolates of P. pacificus cover all four worldwide clades of the species. This extraordinary haplotype diversity suggests multiple independent invasions, most likely in association with different scarab beetles. Together, we establish Réunion as a case study for nematode island biogeography, in which the analysis of nematode population genetics and population dynamics can provide insight into evolutionary and ecological processes. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 170–179.     

Reproductive compatibility and genetic and morphometric variability among populations of the predatory mite,Amblyseius largoensis (Acari: Phytoseiidae), from Indian Ocean Islands and the Americas

The red palm mite (RPM), Raoiella indica Hirst (Acari: Tenuipalpidae), is an invasive phytophagous mite that was recently introduced into The Americas. The predatory mite Amblyseius largoensis Muma (Acari: Phytoseiidae) has been the only natural enemy consistently found in association with RPM. This study aimed to determine if A. largoensis populations from the Indian Ocean Islands (La Réunion and Mauritius) and the Americas (Brazil, Trinidad and Tobago and the USA) consist a taxonomic unit or a group of cryptic species. First, the morphological variability among the A. largoensis populations from these areas was evaluated through morphometric analyses of 36 morphological traits. Then, their genetic variability and phylogenetic relationships were assessed based on two target DNA fragments: the nuclear Internal Transcribed Spacer and the mithochondrial 12S rRNA. Finally, reproductive compatibility of the populations from La Réunion and Roraima, Brazil was evaluated. Morphometric differences between the A. largoensis specimens from La Réunion Island and the Americas were observed, most of them on the length of the setae. Molecular analysis indicated that the A. largoensis populations from the Indian Ocean Islands and the Americas belong to the same taxonomic entity, although to two well defined genetic groups. Crossings involving the A. largoensis populations from La Réunion Island and Roraima, Brazil revealed complete reproductive compatibility between these populations. Information on the morphometric and genetic variability among studied A. largoensis populations can be further exploited in future studies to follow colonization of Indian Ocean Islands populations in the Americas, in the case of field releases.     

Microsatellite DNA markers for the Stable Fly,Stomoxys calcitrans (Diptera: Muscidae)

Eight polymorphic microsatellite markers have been isolated from a microsatellite‐enriched DNA library from the Stable Fly, Stomoxys calcitrans. These loci exhibited four to 15 alleles per locus and an expected heterozygosity ranging from 0.19 to 0.84 in the three populations studied from La Réunion island (Indian Ocean). They should therefore be valuable for studying genetic diversity and population structure. Cross‐species amplification of these eight loci in the closely allied species Stomoxys niger niger was successful for four of these loci, one of which was monomorphic and one of which strongly departed from Hardy–Weinberg expectations.     

Sexual and asexual reproduction of the holothurian Stichopus chloronotus (Echinodermata): a comparison between La Réunion (Indian Ocean) and east Australia (Pacific Ocean)

Summary

Stichopus chloronotus (Brandt, 1835) is one among nine aspidochirotide holothurian species known to reproduce both sexually by broadcast spawning and asexually by transverse fission. New data on the sexual cycle of this species in La Réunion are presented here and information available on sexual and asexual reproduction in this species is summarised. Sexual reproduction on La Réunion shows a distinct seasonality with a main spawning period in the warm season (November-February). The spawning period the Great Barrier Reef appears to be at the same time. Some intriguing deviations from unity in sex-ratio, usually biased towards more male individuals, have been observed in both geographic regions (sex ratio at La Reunion 31:1). New data on the asexual reproduction of this species in La Réunion confirm the high rates of fission. The peak of asexual reproduction in both the Indian and Pacific Ocean was observed in winter (June-July). Thus, asexual reproduction in this species occurs outside the season for sexual reproduction. The rate of asexual reproduction appears to vary between sample locations. However, results of population genetic studies on S. chloronotus (Uthicke et al., 1999; Uthicke et al., 2001) indicated that in most populations investigated a maximum of about 60% of all individuals may be derived from sexual recruitment. Cluster analyses on genetic distances between populations grouped populations within Oceans together, with the exception of one sample from a nearshore reef of the GBR. Although genetic differences between the two regions exist, these are relatively small regarding the large geographic distance. We conclude that asexual reproduction in S. chloronotus is important to maintain local population sizes, but that larval exchange between populations mediated by sexual reproduction is important for colonisation of new areas and to provide connectivity between populations. Here, we present the first synthesis of these phenomena for a holothurian species.     

Unraveling the evolutionary history of the nematode Pristionchus pacificus: from lineage diversification to island colonization

The hermaphroditic nematode Pristionchus pacificus is a model organism with a range of fully developed genetic tools. The species is globally widespread and highly diverse genetically, consisting of four major independent lineages (lineages A, B, C, and D). Despite its young age (~2.1 Ma), volcanic La Réunion Island harbors all four lineages. Ecological and population genetic research studies suggest that this diversity is due to repeated independent island colonizations by P. pacificus. Here, we use model‐based statistical methods to rigorously test hypotheses regarding the evolutionary history of P. pacificus. First, we employ divergence analyses to date diversification events among the four “world” lineages. Next, we examine demographic properties of a subset of four populations (“a”, “b”, “c”, and “d”), present on La Réunion Island. Finally, we use the results of the divergence and demographic analyses to inform a modeling‐based approximate Bayesian computation (ABC) approach, where we test hypotheses about the order and timing of establishment of the Réunion populations. Our dating estimates place the recent common ancestor of P. pacificus lineages at nearly 500,000 generations past. Our demographic analysis supports recent (<150,000 generations) spatial expansion for the island populations, and our ABC approach supports c>a>b>d as the most likely colonization order of the island populations. Collectively, our study comprehensively improves previous inferences about the evolutionary history of P. pacificus.     

Microsatellite loci in the stable fly,Stomoxys niger niger (Diptera: Muscidae) on La Réunion Island

Six polymorphic microsatellite markers have been isolated from a microsatellite‐enriched DNA library from the stable fly, Stomoxys niger niger. These loci exhibited five to 10 alleles per locus and an expected heterozygosity ranging from 0.57 to 0.81 in the studied populations from La Réunion Island (Indian Ocean). They should therefore be useful for the study of genetic diversity and population structure of S. niger. In addition, cross‐species amplification of four of these six loci in the closely allied species Stomoxys calcitrans produced interpretable results, two of which would be useful for population biology studies.     

Tristyly in the endangered Mascarene Island endemic Hugonia serrata (Linaceae)

Tristyly is a rare floral polymorphism known to occur in only five flowering plant families. One unresolved and potential additional case of tristyly concerns the genus Hugonia in the Linaceae. Here we confirm the existence of tristyly in the genus by reporting floral measurements made on Hugonia serrata Lam., an extremely rare species endemic to the Mascarene Islands of La Réunion and Mauritius in the southern Indian Ocean. We conducted an extensive search of all natural habitats on La Réunion Island where the species had been previously reported. Twenty-eight individuals were found, of which nine were in flower. Of the nine flowering individuals five had long-styled flowers with the stigmas placed above the two levels of anthers, three had intermediate length styles with the stigmas placed between the two anther levels, and one had stigmas placed below the two anther levels, i.e., three floral morphs could be identified based on the sequence of stigma and anther positions. Reciprocity and precision indices calculated for sexual organ length in each morph confirm that this variation is well within the range of values observed by previous workers on other tristylous species in other families. Our empirical data confirm the existence of tristyly in the genus Hugonia, thereby raising the number of known families in which tristyly occurs to six. Pollen size, pollen number, and anther length increased slightly with stamen length, but the low number of plants precludes statistical tests of these trends. A bibliographic survey suggests that tristyly may occur in several other species of the genus.     

Natural variation in chemosensation: lessons from an island nematode

All organisms must interact with their environment, responding in behavioral, chemical, and other ways to various stimuli throughout their life cycles. Characterizing traits that directly represent an organism's ability to sense and react to their environment provides useful insight into the evolution of life‐history strategies. One such trait for the nematode Pristionchus pacificus, chemosensation, is involved in navigation to beetle hosts. Essential for the survival of the nematode, chemosensory behavior may be subject to variation as nematodes discriminate among chemical cues to complete their life cycle. We examine this hypothesis using natural isolates of P. pacificus from La Réunion Island. We select strains from a variety of La Réunion beetle hosts and geographic locations and examine their chemoattraction response toward organic compounds, beetle washes, and live beetles. We find that nematodes show significant differences in their response to various chemicals and are able to chemotax to live beetles in a novel assay. Further, strains can discriminate among different cues, showing more similar responses toward beetle washes than to organic compounds in cluster analyses. However, we find that variance in chemoattraction response is not significantly associated with temperature, location, or beetle host. Rather, strains show a more concerted response toward compounds they most likely directly encounter in the wild. We suggest that divergence in odor‐guided behavior in P. pacificus may therefore have an important ecological component.     

Casuarina Invasion Alters Primary Succession on Lava Flows on La Réunion Island

Invasive plants can alter community dynamics and the successional trajectories of ecosystems they colonize. We explore how interactions between disturbance and invasion govern successional trajectories in the case of Casuarina equisetifolia invading lava flows on La Réunion Island. Surveys from 1972 and 1990 were compared with results of a survey in 2012 to detail progression of the invasion over time. General additive models were used to estimate the influence of altitude, distance to putative source of introduction, and lava flow age on the abundance of C. equisetifolia. Based on the predictions, we estimated the likely rate and eventual extent of spread of the species in the area through time. We placed our findings in the context of a conceptual model of successional processes in the area to highlight how the invasion of C. equisetifolia and natural and human‐mediated disturbances are changing natural vegetation dynamics. The extent of invasion by C. equisetifolia has increased twentyfold over the past 40 yr from 110 ha in 1972 to 2373 ha in 2012. Lava flows have facilitated this spread, and in turn C. equisetifolia has started to radically change successional trajectories, increasing the rate of succession sevenfold. The continued spread of this species poses a major threat to the small area of remaining native lowland rain forests on La Réunion Island, which cover <2 percent of their original extent.     

Importance of host oviposition pattern and plant size for the selection of Trichogramma strains to control the diamondback moth

The aim of this work was to select a candidate strain of Trichogrammatidae (Hymenoptera: Chalcidoidea) to control the diamondback moth, Plutella xylostella L. (Lepidoptera: Plutellidae), an important pest of cabbage in Europe. The parasitic efficiency of Trichogramma chilonis Ishii from Japan and Réunion Island, Trichogramma evanescens Westwood from Egypt, Trichogramma ostriniae Pang & Chen from Japan, and Trichogramma semblidis (Aurivillius) from France was studied with sentinel eggs in greenhouse‐grown cauliflower. The percentage of parasitized eggs was measured 1 m from the release point and 3 days after release. Two factors were studied: host‐egg density (three eggs per stem vs. 50 eggs per stem) and plant size (50–100 cm high plants vs. 100–150 cm high plants). Trichogramma evanescens from Egypt achieved the best parasitism. Parasitism efficiency of T. ostriniae from Japan and T. chilonis from Réunion Island was not influenced by egg density. In contrast, parasitism efficiencies of the three other strains were higher at the lower egg density than at the higher egg density. Parasitism efficiency was lower when the plants were bigger, with the exception of T. chilonis from Réunion Island, for which the level of parasitism was not affected by plant size. Thus, T. chilonis from Réunion Island appeared to be the most suitable Trichogramma strain for use in controlling P. xylostella, regardless of the growth stage of the crop. However, T. ostriniae from Japan may also be more suitable for use on cauliflower crops at the early stage of crop development. If potential problems related to the introduction of exotic species are taken into consideration, it is possible to consider the native T. evanescens.     

Eel community structure,fluvial recruitment of <Emphasis Type="Italic">Anguilla marmorata</Emphasis> and indication for a weak local production of spawners from rivers of Réunion and Mauritius islands

Anguillid eels were sampled from permanent rivers in the Réunion and Mauritius islands, western Indian Ocean, with a standardized electrofishing method. A. marmorata was very dominant, corresponding to 91.7 and 90.7% of all the eels collected in Réunion and Mauritius, respectively. Three other species (A. mossambica, A. bicolor bicolor and A. nebulosa labiata) were also present in both islands. A. marmorata showed a strong altitudinal gradient of densities from the lower to upper zones, especially in the younger stages (TL <250 mm), while A. mossambica was only found in the upper zones and A. bicolor bicolor occurred only in the lower zones (A. nebulosa labiata was rare). The eel species composition in freshwaters of both islands is very similar because these two adjoining islands are located in the same trail of drifting marine larvae. Mean estimated eel biomasses were noticeably low (11.1 and 22.2 kg ha⁻¹ in Réunion and Mauritius islands, respectively), especially when compared to those of other tropical insular systems without any eel fishery (Comoros or Polynesia, more than 100 kg ha⁻¹). Nevertheless, the fluvial recruitment of A. marmorata seemed to be regular during the surveyed period, staggering from October to April. The obvious lack of large eels in Mauritius but more significantly in Réunion suggests a high pressure from traditional fishery, and the local reproductive turnover is uncertain. Because sexual maturation seems to occur at a large body size for A. marmorata, as for temperate species, the Réunion and Mauritius rivers may only have a weak contribution to the regional production of spawners. However, the giant mottled eel population in the western Indian Ocean is believed to be panmictic at the regional scale, and may not rely exclusively on these islands’ contribution. A comparison is made with those of freshwater systems in other tropical islands.     

Phylogeography of the introduced species Rattus rattus in the western Indian Ocean,with special emphasis on the colonization history of Madagascar

Aim To describe the phylogeographic patterns of the black rat, Rattus rattus, from islands in the western Indian Ocean where the species has been introduced (Madagascar and the neighbouring islands of Réunion, Mayotte and Grande Comore), in comparison with the postulated source area (India). Location Western Indian Ocean: India, Arabian Peninsula, East Africa and the islands of Madagascar, Réunion, Grande Comore and Mayotte. Methods Mitochondrial DNA (cytochrome b, tRNA and D‐loop, 1762 bp) was sequenced for 71 individuals from 11 countries in the western Indian Ocean. A partial D‐loop (419 bp) was also sequenced for eight populations from Madagascar (97 individuals), which were analysed in addition to six previously published populations from southern Madagascar. Results Haplotypes from India and the Arabian Peninsula occupied a basal position in the phylogenetic tree, whereas those from islands were distributed in different monophyletic clusters: Madagascar grouped with Mayotte, while Réunion and Grand Comore were present in two other separate groups. The only exception was one individual from Madagascar (out of 190) carrying a haplotype that clustered with those from Réunion and South Africa. ‘Isolation with migration’ simulations favoured a model with no recurrent migration between Oman and Madagascar. Mismatch distribution analyses dated the expansion of Malagasy populations on a time‐scale compatible with human colonization history. Higher haplotype diversity and older expansion times were found on the east coast of Madagascar compared with the central highlands. Main conclusions Phylogeographic patterns supported the hypothesis of human‐mediated colonization of R. rattus from source populations in either the native area (India) or anciently colonized regions (the Arabian Peninsula) to islands of the western Indian Ocean. Despite their proximity, each island has a distinct colonization history. Independent colonization events may have occurred simultaneously in Madagascar and Grande Comore, whereas Mayotte would have been colonized from Madagascar. Réunion was colonized independently, presumably from Europe. Malagasy populations may have originated from a single successful colonization event, followed by rapid expansion, first in coastal zones and then in the central highlands. The congruence of the observed phylogeographic pattern with human colonization events and pathways supports the potential relevance of the black rat in tracing human history.     

Novel Cryphonectriaceae from La Réunion and South Africa,and their pathogenicity on <Emphasis Type="Italic">Eucalyptus</Emphasis>

Fungi in the Cryphonectriaceae are important canker pathogens of plants in the Melastomataceae and Myrtaceae (Myrtales). These fungi are known to undergo host jumps or shifts. In this study, fruiting structures resembling those of Cryphonectriaceae were collected and isolated from dying branches of Syzygium cordatum and root collars of Heteropyxis natalensis in South Africa, and from cankers on the bark of Tibouchina grandifolia in La Réunion. A phylogenetic species concept was used to identify the fungi using partial sequences of the large subunit and internal transcribed spacer regions of the nuclear ribosomal DNA, and two regions of the β-tubulin gene. The results revealed a new genus and species in the Cryphonectriaceae from South Africa that is provided with the name Myrtonectria myrtacearum gen. et sp. nov. Two new species of Celoporthe (Cel.) were recognised from La Réunion and these are described as Cel. borbonica sp. nov. and Cel. tibouchinae sp. nov. The new taxa were mildly pathogenic in pathogenicity tests on a clone of Eucalyptus grandis. Similar to other related taxa in the Cryphonectriaceae, they appear to be endophytes and latent pathogens that could threaten Eucalyptus forestry in the future.     

Morphological and plumage colour variation in the Réunion grey white‐eye (Aves: Zosterops borbonicus): assessing the role of selection

The Réunion grey white‐eye (Zosterops borbonicus), a small passerine endemic to the island of Réunion (Mascarene archipelago), constitutes an extraordinary case of phenotypic variation within a bird species, with conspicuous plumage colour differentiation at a microgeographical scale. To understand whether natural selection could explain such variability, we compared patterns of variation in morphological and plumage colour traits within and among populations. To quantify morphological variation, we used measurements obtained by Frank Gill in the 1960s from 239 individuals collected in 60 localities distributed over the entire island of Réunion. To quantify colour variation, we measured the reflectance spectra of plumage patches of 50 males from a subset of Gill's specimens belonging to the five recognized plumage colour variants and used a visual model to project these colours in an avian‐appropriate, tetrachromatic, colour space. We found that variants occupy different regions of the avian colour space and that between‐variant differences for most plumage patches could be discriminated by the birds. Differences in morphology were also detected, but these were, in general, smaller than colour differences. Overall, we found that variation in both plumage colour and morphology among variants is greater than would be expected if genetic drift alone was responsible for phenotypic divergence. As the plumage colour variants correspond to four geographical forms, our results suggest that phenotypic evolution in the Réunion grey white‐eye is at least partly explained by divergent selection in different habitats or regions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 459–473.     

Botryosphaeriaceae associated with Acacia heterophylla (La Réunion) and Acacia koa (Hawaii)

Acacia koa and A. heterophylla are commonly occurring native trees on the Hawaiian Islands and La Réunion, respectively. A recent phylogenetic study suggested that A. heterophylla renders A. koa paraphyletic, and that the former likely arose from the Hawaiian Islands around 1.4 million years ago. An intriguing question is whether their microbiota is similar, although they occur naturally in two very distant geographical locations. In this study, we compared the fungi in the Botryosphaeriaceae isolated from natural populations of A. koa and A. heterophylla. These fungi were chosen because they commonly occur on woody plants and some are important pathogens. They are also known to have been moved globally on asymptomatic plant materials. Isolates were identified based on comparisons of DNA sequence data for the rDNA-ITS, TEF1-α and β-tubulin loci. Ten Botryosphaeriaceae species were identified, of which four species were specific to A. koa from the Hawaiian Islands and five to A. heterophylla in La Réunion. Only one species, Neofusicoccum parvum, which is known to have a wide global distribution, was common to both hosts. The overall results of this study suggest that although A. koa and A. heterophylla share a recent evolutionary history, they have established independent microbiota, at least in terms of the Botryosphaeriaceae.     

The Invasive Woody Weed Ligustrum robustum subsp. walkeri Threatens Native Forests on La Réunion

One of the last primitive island ecosystems in the Indian Ocean has been invaded since 1969 by the Sri Lankan privet, Ligustrum robustum. L. robustum is still spreading in the forests of La Réunion Island, where only 30% of the original vegetation remains, but where 98% of the primary native vegetation of the Mascarene Islands still exists. On Mauritius, where L. robustum was introduced about 1895, it now forms dense, impenetrable thickets, and its presence has been correlated with the inability of native vegetation to re-establish. We assessed the potential impacts of L. robustum invasion on the native ecosystems of La Réunion and identified the factors of invasibility. We determined the degree of invasion in 12 plots of 156 m² and followed native flora and privet recruitment for 3 years in 12 seedling plots of 39 m². The data show that monocultural L. robustum stands now exist in human-disturbed primary forest patches (3.3 individuals/m² and 80% of total individuals) and high seedling densities (0.3–0.6/m²) occur in the least disturbed patches. L. robustum's rapid growth, high shade tolerance and seed production, bird-assisted seed dispersal and high seedling recruitment contribute to its invasiveness in intact forests. The conservation of the original ecosystems of La Réunion depends on the setting up of a long-term and immediate global control strategy. This revised version was published online in July 2006 with corrections to the Cover Date.