The morphology, distribution and conservation implications of introduced rats, Rattus spp. in the granitic Seychelles

There are no native land mammals in the Seychelles archipelago other than bats. Introduced rats have reduced the conservation value of most islands. This paper compares the results of rat‐trapping carried out on eight islands in the granitic Seychelles, between July 1999 and April 2000. Trapping was carried out in both the dry and wet seasons. Three introduced rodent species were caught, including two species of rat (ship rat Rattus rattus Linnaeus and Norway rat R. norvegicus Berkenhout), but only one Rattus species occurred on each island. Both rat species were smaller than European or Asian conspecifics, and there were variations in the size and appearance of rats on different islands. Inter‐island differences in size and pelage colour are discussed.     

Impact of the introduced yellow crazy ant Anoplolepis gracilipes on Bird Island, Seychelles

The introduced yellow crazy ant or long-legged ant Anoplolepis gracilipes was first reported in Seychelles in 1969 and now occurs on at least nine islands in the Central Seychelles. We describe the yellow crazy ant's effects on vegetation and invertebrate communities on one of these, Bird Island; in 2000, Anoplolepis (first reported in 1991) occurred there at densities at least 80 times higher than on other islands in the Central Seychelles. They were associated with high densities of coccid scale insects on foliage, especially of the native tree Pisonia grandis, in some instances causing tree death. Yellow crazy ants on Bird Island also significantly affected invertebrate communities on foliage and on the ground, both in terms of taxonomic composition and the density of specific taxa, apparently causing the local exclusion of some invertebrates.     

Rats are not the only introduced rodents producing ecosystem impacts on islands

In addition to rats, nutria (Myocastor coypus) and the North American beaver (Castor canadensis) have certainly caused damage at an ecosystem level when introduced to islands, in both cases primarily by ecosystem engineering. Of other introduced rodents successfully established on islands, the gray squirrel (Sciurus carolinensis) may be in the process of damaging entire forest ecosystems, particularly by bark-stripping. Though introduced muskrats (Ondatra zibethicus) have had ecosystem-level impacts in continental Europe, their impact on islands worldwide to which they have been introduced has been very limited. The North American red squirrel (Tamiasciurus hudsonicus) and Barbary ground squirrel (Atlantoxerus getulus) have each had substantial impacts when introduced to particular islands, but for neither species have these impacts yet been demonstrated to spread through an entire ecosystem. Introduced house mice (Mus musculus) may well generate ecosystem impacts on remote islands lacking rats, and it is possible that explosions of house mice on islands after rat eradication, a common occurrence, will lead in some instances to ecosystem impacts.     

Evolution ofPeperomia (Piperaceae) in the Juan Fernandez Islands,Chile

Four species ofPeperomia (Piperaceae) occur in the Juan Fernandez Islands, Chile:P. berteroana, P. margaritifera, P. skottsbergii, andP. fernandeziana. The last species is found also in continental Chile, whereas the other three are endemic to the archipelago.Peperomia margaritifera is found only on the older island of Masatierra, whereasP. skottsbergii is confined to the younger island of Masafuera, andP. berteroana occurs on both islands. Phenetic analyses of mainland taxa suggest thatP. fernandeziana belongs to subg.Sphaerocarpidium whereas the endemic taxa form their own subg.Tildenidium connecting to subg.Tildenia. Cladistic analyses indicate thatP. margaritifera is the most primitive species in the archipelago and thatP. berteroana is the most derived, especially patristically. Chromosomally, the four species are all n = 22, which may be tetraploid on a base of x = 11. Sulfated flavones occur only inP. berteroana andP. skottsbergii, which are otherwise unknown for the family. Dispersal of propagules to the islands from the continent and between islands is believed to have been accomplished by birds.     

Snails of the genus Pachnodus (Mollusca; Gastropoda; Enidae): their origins and evolution

Aim The phylogeny of the enid land-snail genus Pachnodus was determined in order to provide information on biogeographical patterns within the granitic Seychelles islands. Location The genus Pachnodus is endemic to the granitic islands of Seychelles (Mahé, Silhouette, Praslin, La Digue and Fregate). Methods Phylogeny was determined using a cladistic analysis of nineteen shell and soft-body anatomy characters. The outgroup used was the central-east African genus Cerastus. Results The proposed phylogeny indicates that the genus divided into two distinct subgenera early in its history. Subsequent speciation occurred in parallel in the two subgenera, resulting in several islands supporting at least two distinct species representing the two subgenera. Main conclusions The pattern of speciation is largely explicable by vicariance as a result of sea-level rises, followed by habitat specialisation and further speciation. The pattern is in contrast to previously published scenarios for other taxa in the islands and indicates significant differences between evolutionary and habitat factors in the biogeography of the Seychelles fauna     

On some rust fungi (Uredinales) collected in an Acacia koa–Metrosideros polymorpha woodland, Mauna Loa Road, Big Island, Hawaii

Nine species of rust fungi (Uredinales) were found during a 1-day field study in an Acacia koa–Metrosideros polymorpha woodland in Volcanoes National Park on Big Island, Hawaii. Two species, both found on Acacia koa, are native (endemic) rusts whereas 7 species are nonnative on nonnative hosts, highlighting the high proportion of introduced species in the Hawaiian flora even in more or less natural habitats. One species, Uromyces linearis on Panicum repens, constitutes the first record of this rust for the Hawaiian archipelago, bringing the total to 93 species on the islands, 70 (75.3%) of which are introduced. The species records are annotated with emphasis on the geographic origin of each taxon. In addition, the study has led to the molecular reevaluation of the genus Racospermyces, indicating that it is synonymous with Endoraecium, and six new combinations are proposed for the species previously placed in Racospermyces. The high number of introduced species in Hawaii coupled with the paucity of native species when compared to other global regions is discussed.     

Song dialects on the Atlantic islands: goldcrests of the Azores ( Regulus regulus azoricus, R. r. sanctae-mariae, R. r. inermis)

Territorial songs in island populations of songbirds are often highly divergent from those of neighbouring continental relatives. This is shown for the three goldcrest subspecies (Regulus regulus azoricus, R. r. sanctae-mariae and R. r. inermis) endemic on six islands of the Azorean archipelago. All investigated populations display a high intra- and inter-individual acoustic variation. On each island, up to six different song types have been found; and a single male sings up to three types. In contrast, all northwestern European populations of R. r. regulus and R. r. anglorum share only a single song type. In playback experiments, none of 18 tested dialect songs of Azorean goldcrests evoked notable territorial reaction in German and Czech goldcrest males (ssp. regulus). Two differing dialect groups of the goldcrest can be distinguished on the Azores. Populations of the eastern islands, São Miguel and Santa Maria, share common song types which are not found on the islands of the central and western groups. Dialect repertoires on the westernmost islands, Flores, Faial and Pico, are dominated by a different song type. In the geologically younger western crater of São Miguel, both western and eastern song types coexist. Acoustic similarities to a population from neighbouring Terceira suggest the western part of São Miguel as the origin for the westward expansion of R. regulus on the Azores.     

Presence of the Chytrid Fungus Batrachochytrium dendrobatidis in Populations of the Critically Endangered Frog Mannophryne olmonae in Tobago, West Indies

The emerging infectious disease chytridiomycosis is prevalent in Central and South America, and has caused catastrophic declines of amphibian populations in the Neotropics. The responsible organism, Batrachochytrium dendrobatidis, has been recorded on three West Indian islands, but the whole of the Caribbean region is predicted to offer a suitable environment for the disease. Monitoring the spread of chytridiomycosis is thus a priority in this region, which has exceptionally high levels of amphibian endemism. PCR analysis of 124 amphibian skin swabs in Tobago (Republic of Trinidad and Tobago) demonstrated the presence of B. dendrobatidis in three widely separated populations of the frog Mannophryne olmonae, which is listed as Critically Endangered on the basis of recent population declines. Chytridiomycosis is presently endemic in this species, with a prevalence of about 20% and no associated clinical disease. Increased susceptibility to chytridiomycosis from climate change is unlikely in amphibian populations in Tobago, as this island does not have high montane environments, but remains a possibility in the sister island of Trinidad. Preventing the spread of chytridiomycosis within and between these and other Caribbean islands should be a major goal of practical conservation measures for amphibians in the region.     

Hybridization occurs between Drosophila simulans and D. sechellia in the Seychelles archipelago

Drosophila simulans and D. sechellia are sister species that serve as a model to study the evolution of reproductive isolation. While D. simulans is a human commensal that has spread all over the world, D. sechellia is restricted to the Seychelles archipelago and is found to breed exclusively on the toxic fruit of Morinda citrifolia. We surveyed the relative frequency of males from these two species in a variety of substrates found on five islands of the Seychelles archipelago. We sampled different fruits and found that putative D. simulans can be found in a variety of substrates, including, surprisingly, M. citrifolia. Putative D. sechellia was found preferentially on M. citrifolia fruits, but a small proportion was found in other substrates. Our survey also shows the existence of putative hybrid males in areas where D. simulans is present in Seychelles. The results from this field survey support the hypothesis of current interbreeding between these species in the central islands of Seychelles and open the possibility for fine measurements of admixture between these two Drosophila species to be made.     

Population Genetic Structure and Conservation of the Galápagos Petrel (Pterodroma phaeopygia)

The Galápagos petrel (Pterodroma phaeopygia) is endemic to the Galápagos archipelago, where it is known to breed only on five islands. The species has been listed as critically endangered due to habitat deterioration and predation by introduced mammals. Significant morphological and behavioural differences among petrels nesting on different islands suggest that island populations may differ genetically. Furthermore, nesting phenology suggests that genetically differentiated seasonal populations may exist within at least one island. We analysed variation in six microsatellite loci and part of the mitochondrial ATPase 6/8 gene in 206 Galápagos petrels sampled from all five islands. No evidence of genetic structuring within islands was found, although statistical power was low. In contrast, significant differences occurred among island populations. For the microsatellite loci, private alleles occurred at all islands, sometimes at high frequency; global and pairwise estimates of genetic differentiation were all statistically significant; Bayesian analysis of genotypes frequencies provided strong support for three genetic populations; and most estimates of migration between populations did not differ significantly from zero. Only two ATPase haplotypes were found, but the geographic distribution of haplotypes indicated significant differentiation among populations. For conservation purposes, populations from Floreana, Santa Cruz, San Cristóbal and Santiago should be regarded as separate genetic management units. Birds from Isabela appear to be derived recently from the Santiago population, and the population on San Cristóbal appears to be a mixture of birds from other populations. However, considering ecological and behavioural differences among birds from different islands, we recommend that all five populations be protected.     

Microsatellite variation suggests a recent fine-scale population structure of <Emphasis Type="Italic">Drosophila sechellia</Emphasis>, a species endemic of the Seychelles archipelago

Drosophila sechellia is closely related to the cosmopolitan and widespread model species, D. simulans. This species, endemic to the Seychelles archipelago, is specialized on the fruits of Morinda citrifolia, and harbours the lowest overall genetic diversity compared to other species of Drosophila. This low diversity is associated with a small population size. In addition, no obvious population structure has been evidenced so far across islands of the Seychelles archipelago. Here, a microsatellite panel of 17 loci in ten populations from nine islands of the Seychelles was used to assess the effect of the D. sechellia’s fragmented distribution on the fine-scale population genetic structure, the migration pattern, as well as on the demography of the species. Contrary to previous results, also based on microsatellites, no evidence for population contraction in D. sechellia was found. The results confirm previous studies based on gene sequence polymorphism that showed a long-term stable population size for this species. Interestingly, a pattern of Isolation By Distance which had not been described yet in D. sechellia was found, with evidence of first-generation migrants between some neighbouring islands. Bayesian structuring algorithm results were consistent with a split of D. sechellia into two main groups of populations: Silhouette/Mahé versus all the other islands. Thus, microsatellites suggest that variability in D. sechellia is most likely explained by local genetic exchanges between neighbouring islands that have recently resulted in slight differentiation of the two largest island populations from all the others.     

Successful spread of a biocontrol agent reveals a biosecurity failure: elucidating long distance invasion pathways for <Emphasis Type="Italic">Gonatocerus ashmeadi</Emphasis> in French Polynesia

Invasive species are generally detected in new ecosystems long after their first arrival, making it difficult to elucidate pathways leading to successful invasion. In this study, the dispersal of a classical biological control agent, the mymarid egg parasitoid Gonatocerus ashmeadi, was monitored across ten islands in three major island groups in French Polynesia from the exact moment of its introduction into Tahiti to combat the invasive pest Homalodisca vitripennis. Within 10 months, the parasitoid spread quickly from Tahiti to widely separated islands (up to 1,400 km from Tahiti); presumably through the transportation of plant material containing parasitized H. vitripennis eggs. Gonatocerus ashmeadi thus functioned as a “biomarker”, providing an informal audit of the effectiveness of inter-island quarantine measures designed to curb the accidental spread of noxious organisms. Survey results suggest that invasive organisms, like deliberately released biological control agents, can be unintentionally and rapidly transmitted across vast distances by humans. Furthermore, even remote islands appear to experience relentless pressure from invasive propagules associated with human travel. Implications of survey work documenting the spread and impact of G. ashmeadi are discussed within the context of biological control programs, non-target impacts, and biosecurity initiatives. Handling Editor: Dirk Babendreier.     

Restricted plant species on sub-Antarctic Macquarie and Heard Islands

The recent distributional history of two Macquarie Island vascular plant species, Carex trifida, Poa litorosa, and the Heard Island vascular plant, Ranunculus crassipes is examined. C. trifida is known from only one small population on the north west coast of Macquarie Island. Four populations of P. litorosa were first recorded in the 1980s; we believe however, that it was first observed, but misidentified in the 1950s. R. crassipes was first discovered on Heard Island in the late 1980s. We argue that all three species are indigenous and arrived on their respective islands within the last 200 years by natural processes, most likely from warmer neighbouring islands, where these species have more extensive distributions. There have been small-scale changes in distribution of all species, mainly expansion. Further expansion of all three species is expected as a response to warming climate. Feral rabbit grazing is having a confounding negative influence on populations of P. litorosa.     

Consistent performance of invasive plant species within and among islands of the Mediterranean basin

Since the success of an invasive species depends not only upon its intrinsic traits but also on particular characteristics of the recipient habitat, assessing the performance of an invader across habitats provides a more realistic analysis of risk. Such an analysis will not only provide insights into the traits related to invasiveness, but also the habitat characteristics that underpin vulnerability to invasion that, taken together, will facilitate the selection of management strategies to mitigate the invader’s effect. In the present study, we considered the Mediterranean basin islands as an excellent study region to test how the same invasive species perform in different habitats within a single island, and to scale up differences among islands with similar climate. We tested how the performance of three widespread plant invaders with clonal growth but contrasting life-history traits, a deciduous tree Ailanthus altissima, a succulent subshrub Carpobrotus spp., and an annual geophyte Oxalis pes-caprae, varied depending upon the species identity, habitat, and invaded island. The environmental parameters considered were habitat type, elevation, species diversity in the invaded plot, and several soil traits (% C, % N, C/N, pH, and relative humidity). The study documents that the performance of these three important and widespread plant invaders is dependent mainly on species identity, and less upon the invaded island’s general features. Likewise, differences in performance among habitats were only significant in the case of Ailanthus, whereas Carpobrotus and Oxalis appear to perform equally well in different environments. Ailanthus thus appears to have a broader spectrum of invasiveness, being able to invade a larger number of habitat types. On the contrary, Carpobrotus spp. have not yet invaded habitats different from those where the species have been originally introduced and where they are still commonly spread by humans. Oxalis distribution is mainly related to agricultural activities and disturbed sites, and the total area infested by this geophyte may be more reflection of the extent of suitable habitats than of invasiveness or ecological impact. Our results confirm the potential for these species to significantly alter the functioning of ecosystems in the Mediterranean islands and highlight the risk to other islands not yet invaded.     

Invasion of the Hawaiian Islands by a parasite infecting imperiled stream fishes

Points of origin and pathways of spread are often poorly understood for introduced parasites that drive disease emergence in imperiled native species. Co‐introduction of parasites with non‐native hosts is of particular concern in remote areas like the Hawaiian Islands, where the introduced nematode Camallanus cotti has become the most prevalent parasite of at‐risk native stream fishes. In this study, we evaluated the prevailing hypothesis that C. cotti entered the Hawaiian Islands with poeciliid fishes from the Americas, and spread by translocation of poeciliid hosts across the archipelago for mosquito control. We also considered the alternative hypothesis of multiple independent co‐introductions with host fishes originating from Asia. We inferred conduits of introduction and spread of C. cotti across the archipelago from geographic patterns of mtDNA sequence variation and allelic variation across 11 newly developed microsatellite markers. The distribution of haplotypes suggests that C. cotti spread across the archipelago following an initial introduction on O'ahu. Approximate Bayesian Computation modeling and allelic variation also indicate that O'ahu is the most likely location of introduction, from which C. cotti dispersed to Maui followed by spread to the other islands in the archipelago. Evidence of significant genetic structure across islands indicates that contemporary dispersal is limited. Our findings parallel historical records of non‐native poeciliid introductions and suggest that remediating invasion hotspots could reduce the risk of infection in native stream fishes, which illustrates how inferences on parasite co‐introductions can improve conservation efforts by guiding responses to emerging infectious disease in species of concern.     

Modelling the distribution and interaction of introduced rodents on New Zealand offshore islands

Aim To establish the factors that correlate with the distribution of the four most commonly introduced rodent species on New Zealand offshore islands — ship rat (Rattus rattus), Norway rat (R. norvegicus), Pacific rat or kiore (R. exulans) and house mouse (Mus musculus) — and examine if these distributions are interactive at the archipelago scale. Location The 297 offshore islands of the New Zealand archipelago (latitude: 34° S to 47° S; longitude: 166° E to 179° E). Methods Data on the distribution of all four introduced rodent species and the characteristics of New Zealand offshore islands were collated from published surveys and maps. The distribution of individual rodent species was regressed on island characteristics using a logistic generalized linear model. Interactions were examined by including the distributions of other rodent species as predictors in models. Results All four rodent species appear to be limited by a variety of factors, which differ between species in both number and type. The distribution of ship rats is limited by the most factors, reflecting the extent of its distribution across the archipelago. The distribution of mice is the least explicative. Only the three rat species interacted in their distribution. The distribution of kiore on offshore islands is significantly negatively related to that of ship rats and to a lesser extent Norway rats. The distribution of mice did not appear affected in any way by the number of other rodent species on an island. Main conclusions Differences in competitive ability and dispersal allow all four species to inhabit the New Zealand archipelago. Kiore distribution appears to be most limited by ship rat (and to a lesser extent Norway rat) distribution. The distribution of kiore was not found to interact with the distribution of mice on offshore islands, as has been suggested by others. The distribution of mice on offshore islands was difficult to model, which highlights the difficulties in managing this species. Overall the results offer valuable insights for management methods to assist preventing the invasion of offshore islands.     

The role of exotic plants in the invasion of Seychelles by the polyphagous insect <Emphasis Type="Italic">Aleurodicus dispersus</Emphasis>: a phylogenetically controlled analysis

The accidental introduction of the spiralling whitefly, Aleurodicus dispersus Russell (Homoptera: Aleyrodidae) to Seychelles in late 2003 is exploited during early 2005 to study interactions between A. dispersus, native and exotic host plants and their associated arthropod fauna. The numbers of A. dispersus egg spirals and pupae, predator and herbivore taxa were recorded for eight related native/exotic pairs of host plants found on Mahé, the largest island in Seychelles. Our data revealed no significant difference in herbivore density (excluding A. dispersus) between related native and exotic plants, which suggests that the exotic plants do not benefit from ‘enemy release’. There were also no differences in predator density, or combined species richness between native and exotic plants. Together these data suggest that ‘biotic resistance’ to invasion is also unlikely. Despite the apparent lack of differences in community structure significantly fewer A. dispersus egg spirals and pupae were found on the native plants than on the exotic plants. Additional data on A. dispersus density were collected on Cousin Island, a managed nature reserve in which exotic plants are carefully controlled. Significantly higher densities of A. dispersus were observed on Mahé, where exotic plants are abundant, than on Cousin. These data suggest that the rapid invasion of Seychelles by A. dispersus may largely be due to the high proportion of plant species that are both exotic and hosts of A. dispersus; no support was found for either the ‘enemy release’ or the ‘biotic resistance’ hypotheses.     

ADAPTIVE RADIATION OF DAY-GECKOS (PHELSUMA) IN THE SEYCHELLES ARCHIPELAGO: A PHYLOGENETIC ANALYSIS

A phylogenetic analysis using characters derived from mitochondrial DNA was used to show that the species of Phelsuma in the Seychelles Islands represent a single, monophyletic lineage that has diversified as a result of both historical and ecological factors. In the distant past, the Seychelles archipelago was physically invaded by a single species of Phesluma. Separate eustatic sea level changes likely led first to allopatric speciation and then to the secondary contact of these sister species. Differences in the relative timing of the secondary contact between island groups resulted in P. sundbergi evolving an intermediate body size in the group of islands associated with Mahé and a large body size, while sympatric with P. astriata, in the group of islands associated with Praslin. Ecological information was used to support the conclusion that the actual evolutionary mechanism for the body size shift was a response to frequency dependent natural selection of P. sundbergi in single-species and two-species competitive regimes.     

Distribution of clubroot disease of a cruciferous weed, Cardamine flexuosa, in major isolated islands, Hokkaido and Okinawa in Japan

To investigate the distribution of clubroot of a cruciferous weed, Cardamine flexuosa, caused by Plasmodiophora brassicae, field surveys were conducted in Hokkaido, Aomori, and Okinawa, and major isolated islands in Japan during 1993–2004. The disease was newly recorded in Aomori and nine islands in five different prefectures, including Sado (Niigata), Oki (Shimane), Mishima (Yamaguchi), Tsushima, Iki and Goto (Nagasaki), and Koshiki, Yakushima, and Tanegashima (Kagoshima). The diseased plants were not found in Hokkaido and Okinawa (islands of Okinawa, Kumejima, Ishigaki, Iriomote, and Kohama). However, inoculation tests showed that most C. flexuosa collected from Hokkaido and Okinawa included many susceptible plants. The result suggests that resistance of the plants is not the reason that the disease was not found in these areas. An erratum to this article is available at .     

The genetics of adaptation in Drosophila sechellia

Drosophila sechellia is an island endemic of the Seychelles. After its geographic isolation on these islands, D. sechellia evolved into a host specialist on the fruit of Morinda citrifolia – a fruit often noxious and repulsive to Drosophila. Specialization on M. citrifolia required the evolution of a suite of adaptations, including resistance to and preference for some of the toxins found in this fruit. Several of these adaptive traits have been studied genetically. Here, I summarize what is known about the genetics of these traits and briefly describe the ecological and geographical context that shaped the evolution of these characters. The data from D. sechellia suggest that adaptations are not as genetically complex as historically thought, although almost all of the adaptations of D. sechellia involve several genes.