Genetic Connectivity of the Moth Pollinated Tree Glionnetia sericea in a Highly Fragmented Habitat

Long-distance gene flow is thought to be one prerequisite for the persistence of plant species in fragmented environments. Human influences have led to severe fragmentation of native habitats in the Seychelles islands, with many species surviving only in small and isolated populations. The endangered Seychelles endemic tree Glionnetia sericea is restricted to altitudes between 450 m and 900 m where the native forest vegetation has been largely lost and replaced with exotic invasives over the last 200 years. This study explores the genetic and ecological consequences of population fragmentation in this species by analysing patterns of genetic diversity in a sample of adults, juveniles and seeds, and by using controlled pollination experiments. Our results show no decrease in genetic diversity and no increase in genetic structuring from adult to juvenile cohorts. Despite significant inbreeding in some populations, there is no evidence of higher inbreeding in juvenile cohorts relative to adults. A Bayesian structure analysis and a tentative paternity analysis indicate extensive historical and contemporary gene flow among remnant populations. Pollination experiments and a paternity analysis show that Glionnetia sericea is self-compatible. Nevertheless, outcrossing is present with 7% of mating events resulting from pollen transfer between populations. Artificial pollination provided no evidence for pollen limitation in isolated populations. The highly mobile and specialized hawkmoth pollinators (Agrius convolvuli and Cenophodes tamsi; Sphingidae) appear to promote extensive gene flow, thus mitigating the potential negative ecological and genetic effects of habitat fragmentation in this species. We conclude that contemporary gene flow is sufficient to maintain genetic connectivity in this rare and restricted Seychelles endemic, in contrast to other island endemic tree species with limited contemporary gene flow.     

Insights into the feeding ecology of the Seychelles Black Parrot Coracopsis barklyi using two monitoring approaches

Feeding ecology is an important factor for the survival of a species and knowledge of its parameters is a prerequisite for successful conservation work. In this study we describe the feeding ecology of the endemic Seychelles Black Parrot Coracopsis barklyi on Praslin, Seychelles, the only island on which this parrot is resident. We compared two methods to evaluate feeding choices: incidental observations and feeding walks on 25 transects in all habitat types. Black parrots fed on 46 different species, bringing the total number of known food plants to 53 species. They predominantly consumed endemic and native species (58% of observed feeding bouts), mainly their fruit pulp (in 68% of feeding bouts), followed by buds (15%) and seeds (37%) with occasional observations of leaves, bark and scale insects. The incidental method rendered many more observed bouts than the transect approach and the ratios of consumed species differed between methods but the transect results are regarded as more representative. The incidental method is not suitable for quantitative conclusions but complements the transect method, providing information about rarely occurring feeding events.     

Diversity and conservation of forest-floor arthropods on a small Seychelles island

Cousine Island, Seychelles, is of major conservation significance as it is in a biodiversity hotspot. Furthermore, it is relatively pristine, and is apparently the only tropical island over 20 ha with no alien invasive mammals. This study focuses on the island's log and litter arthropods, which were sampled by extraction methods from the dominant species, Pisonia grandis, Ficus spp. and Cocos nucifera. Stage of decomposition, and forest type in which the logs occurred, both significantly influenced the composition and structure of the arthropod assemblages. Young logs were significantly richer in species than older logs, possibly because they had the most resources and microhabitats. There were some significant changes in arthropod species richness, composition and abundance between species of young logs, but not old ones, because as logs decomposed, arthropod assemblages converged. Nevertheless, each old log species had some arthropod species not present in other log species, which has important implications for conservation. Arthropod assemblages in woody litter varied according to the forest type in which they occurred, and were different from those in logs in the same forest type. Cousine Island arthropod species richness, both in logs and litter, was comparable to figures from other tropical areas. As the logs, especially P. grandis, are home to many Seychelles endemic species, their conservation is essential. Furthermore, as the arthropods are also the main food of certain threatened Seychelles vertebrates, their conservation also underpins a food chain on this unique tropical island.     

Ectomycorrhizal fungi of the Seychelles: diversity patterns and host shifts from the native Vateriopsis seychellarum (Dipterocarpaceae) and Intsia bijuga (Caesalpiniaceae) to the introduced Eucalyptus robusta (Myrtaceae), but not Pinus caribea (Pinaceae)

Ectomycorrhizal (ECM) fungi form highly diverse communities in temperate forests, but little is known about their community ecology in tropical ecosystems. Using anatomotyping and rDNA sequencing, ECM fungi were identified on root tips of the introduced Eucalyptus robusta and Pinus caribea as well as the endemic Vateriopsis seychellarum and indigenous Intsia bijuga in the Seychelles. Sequencing revealed 30 species of ECM fungi on root tips of V. seychellarum and I. bijuga, with three species overlapping. Eucalyptus robusta shared five of these taxa, whereas P. caribea hosted three unique species of ECM fungi that were likely cointroduced with containerized seedlings. The thelephoroid (including the anamorphic genus Riessiella), euagaric, boletoid and hymenochaetoid clades of basidiomycetes dominated the ECM fungal community of native trees. Two species of Annulatascaceae (Sordariales, Ascomycota) were identified and described as ECM symbionts of V. seychellarum. The low diversity of native ECM fungi is attributed to deforestation and long-term isolation of the Seychelles. Native ECM fungi associate with exotic eucalypts, whereas cointroduced ECM fungi persist in pine plantations for decades.     

Are the native giant tortoises from the Seychelles really extinct? A genetic perspective based on mtDNA and microsatellite data

The extinction of the giant tortoises of the Seychelles Archipelago has long been suspected but is not beyond doubt. A recent morphological study of the giant tortoises of the western Indian Ocean concluded that specimens of two native Seychelles species survive in captivity today alongside giant tortoises of Aldabra, which are numerous in zoos as well as in the wild. This claim has been controversial because some of the morphological characters used to identify these species, several measures of carapace morphology, are reputed to be quite sensitive to captive conditions. Nonetheless, the potential survival of giant tortoise species previously thought extinct presents an exciting scenario for conservation. We used mitochondrial DNA sequences and nuclear microsatellites to examine the validity of the rediscovered species of Seychelles giant tortoises. Our results indicate that the morphotypes suspected to represent Seychelles species do not show levels of variation and genetic structuring consistent with long periods of reproductive isolation. We found no variation in the mitochondrial control region among 55 individuals examined and no genetic structuring in eight microsatellite loci, pointing to the survival of just a single lineage of Indian Ocean tortoises.     

Variation in Herbivore Damage to Invasive and Native Woody Plant Species in Open Forest Vegetation on Mahé, Seychelles

Enemy release of introduced plants and variation in herbivore pressure in relation to community diversity are presently discussed as factors that affect plant species invasiveness or habitat invasibility. So far few data are available on this topic and the results are inconclusive. We compared leaf herbivory between native and invasive woody plants on Mahé, the main island of the tropical Seychelles. We further investigated variation in leaf herbivory on three abundant invasive species along an altitudinal gradient (50–550 m a.s.l.). The median percentage of leaves affected by herbivores was significantly higher in native species (50%) than in invasive species (27%). In addition, the species suffering from the highest leaf area loss were native to the Seychelles. These results are consistent with the enemy release hypothesis (ERH). While the invasive species showed significant and mostly consistent variation in the amount of leaf damage between sites, this variation was not related to general altitudinal trends in diversity but rather to local variation in habitat structure and diversity. Our results indicate that in the Seychelles invasive woody plants profit from herbivore release relative to the native species and that the amount of herbivory, and therefore its effect on species invasiveness or habitat invasibility, may be dependent on local community structure and composition.     

Why Seychelles Warblers fail to recolonize nearby islands: unwilling or unable to fly there?

The Seychelles Warbler Acrocephalus sechellensis is a rare island endemic which, from 1920 to 1988, occurred only on Cousin Island (29 ha) in the Seychelles. Despite the saturated nature of this population and the possibility of obtaining higher reproductive success on new nearby islands, inter-island dispersal by Seychelles Warblers is extremely rare (0.10%). We test the hypothesis that Seychelles Warblers show an adaptation typical for island birds: a low-cost reduced-size flight apparatus. We compared the anatomy of the flight apparatus (wing shape, wing loading, skeletal parts and musculature) of Seychelles Warblers with that of three closely related migratory Acrocephalus species (Eurasian Reed Warbler A. scirpaceus , Australian Reed Warbler A. australis and African Reed Warbler A. baeticatus ). Seychelles Warblers do not differ from the migratory warblers in pectoral mass and skeletal attachment area relative to body mass, wing shape and wing loading. Seychelles Warblers show the morphological structures required for sustained flight, but may have the behavioural reluctance to cross what they may regard as extensive bodies of water.     

Trichomonad infection in endemic and introduced columbids in the Seychelles

Island endemic avifaunas face many threats, including the now well-documented impacts of pathogens. The impacts of pathogens on the endemic Seychelles avifauna, however, have been little studied. The protozoan parasite Trichomonas gallinae has been shown to reduce survival and reproductive success of the endemic Pink Pigeon Columba mayeri on the nearby island of Mauritius. I investigated trichomonad infection prevalence and pathogenicity in endemic Seychelles Blue Pigeons, Alectroenas pulcherrima, and two introduced species of columbid, the Madagascar Turtle-dove, Streptopelia picturata, and the Barred Ground Dove, Geopelia striata, on the Seychelles island of Mahé during September-October 2007. I asked whether: 1) trichomonad infections occur in these species; 2) prevalence varies among species; and 3) birds show any signs of pathogenicity consistent with tricho-monosis. I use the results to assess the potential threat of this pathogen to A. pulcherrima. All three species were infected with trichomonads, and the overall prevalence was 27.5%. Alectroenas pulcherrima had higher prevalence (47.1%) than the two introduced species combined (24.3%). No infected individuals showed any signs of disease. These findings suggest that trichomonad parasites should be considered as a potential disease threat to the A. pulcherrima population.     

Population estimates and distribution changes of landbirds on Silhouette Island,Seychelles

Cresswell, W., Irwin, M., Mee, A., Mellanby, R., McKean, M. & Milne, L. 1997. Population estimates and distribution changes of landbirds on Silhouette Island, Seychelles. Ostrich 68 (2–4): 50–57.

The density of landbird species within different habitats, and presence within 4 ha squares, on Silhouette Island were recorded in August 1996, repeating a previous survey (January 1979). Densities of the four most common species were calculated from point counts: Seychelles Sunbirds occurred at 2–5 birds/ha, Seychelles Bulbuls 0.5–4 birds/ha, Indian Mynahs 0.2–2 birds/ha and Seychelles Blue Pigeons 0.3 birds/ha depending on habitat. Population estimates were: Seychelles Sunbirds 5600, Seychelles Bulbuls 4000, Indian Mynahs 1100, Seychelles Blue Pigeons 600, and Seychelles Kestrel, approximately 15 pairs. The distributions of non-endemic Indian Mynahs, Barred Ground Doves and Madagascar Fodies were smaller in 1996 compared to 1979, while those of endemic Seychelles Bulbuls and Sunbirds were larger. Mynahs were more likely to occur in squares containing human activity, but habitat variables rather than human effects predicted the presence or absence of endemics. Mynahs were more likely to occur in a square, controlling for habitat variables, if Bulbuls were absent, and vice versa. There may be major seasonal changes in the relative distribution of introduced and endemic species. Alternatively, introduced species may have actually declined, probably because of a decrease in human population between the two surveys; endemics may then have increased because of reduced competition.     

Number of endemic and native plant species in the Galápagos Archipelago in relation to geographical parameters

By simple and multiple regression analyses we investigate updated species numbers of endemic and native vascular plants and seed plants in the Galápagos Archipelago in relation to geographical parameters. We find that the best models to describe species numbers are regression models with log-transformed species numbers as dependent and log-transformed modified area (i.e. area not covered with barren lava) as an independent variable. This holds both for total species number, for native species number, for endemic species number and for total number of seed plants as well as number of endemic seed plants. For the ratio between endemic and native species, modified area is also the major significant variable, but with a negative regression slope.
Multiple regression models show that some isolation measures are significant contributors and may explain some of the residual variation, but their contribution to total explained variation is in general small.
The results show that the species area relationships are different for native and endemic species. This is discussed in relation to classical island biogeographical models, and the concepts of radiative speciation.     

Possible roles of introduced plants for native vertebrate conservation: the case of Madagascar

Restoration approaches rely on native plants; yet in some situations, natural vegetation may not grow fast enough to prevent the fragmentation of original vegetation and the consequent negative impacts on fauna. In this context, some introduced plants may grow faster and provide more food than native species, and they may also contribute to human livelihood. We investigate to what extent introduced plant species (1) can serve as habitat and food for endemic vertebrates and (2) provide benefits to local people. We address this question in Madagascar, characterized by high degrees of endemism, long histories of coevolution between endemic species, highly fragmented forests, and a high reliance of the rural population on natural resources. A literature search for interactions between endemic fauna and introduced flora revealed that 100 of 1,379 introduced species recorded for Madagascar are used by endemic vertebrates. They provide food mainly for primates, flying foxes, and birds, and habitat for all terrestrial vertebrate groups. One hundred vertebrate species were reported to use introduced plants, many of which are fast growing and are useful for populations. Although these introduced plants should be approached with caution due to their potentially invasive behavior, many introduced plants can provide services for the native fauna and for humans. For example, trees can provide an interim solution to secure the survival of endemic fauna that otherwise would be lost due to fragmentation effects. These plants could bridge the time lag until native forest regeneration or restoration with native trees will have become effective.     

Restoration of a tropical island: Cousine Island,Seychelles

Tropical island species and ecosystems are threatened worldwide as a result of increasing human pressure. Yet some of these islands also lend themselves to restoration, as they are physically defined units that can be given focused attention, as long as resources are available and clear conservation targets are set. Cousine Island, Seychelles, is a tropical island that has received such intensive restoration. From a highly degraded island in the 1960s, the island has now been restored to what is believed to be a semblance of the natural state. All alien vertebrates have been eradicated, as have 25 invasive alien plants. Cultivated plants are now confined to one small section of the island. Poaching of nesting marine turtles has been stopped, leading to an increase in turtle breeding numbers. The shearwater population has increased in size with poaching activities under control. The Sooty tern has also returned to the island to breed. The coastal plain has been restored with over 2,500 indigenous shrubs and trees, which have now grown into a forest carpet. There are strict quarantine procedures on the island, keeping it free of rats, mice, various alien invertebrates and potentially invasive alien plants. Three threatened Seychelles endemic land birds (Seychelles warbler, Seychelles magpie robin and Seychelles white-eye) have been introduced and are thriving, with these introductions contributing to both the magpie robin and the white-eye being downgraded from CR to EN (the warbler remains at VU). Ecotourism, and nature conservation for the local inhabitants, have been introduced in a way that does not reduce the improved compositional, structural and functional biodiversity of the island. The result of the restoration effort appears to be sustainable in the long term, although challenges still remain, especially with regards to adequate clean water and a non-polluting power supply on the island. Cousine is thus paving the way in the art and science of tropical island restoration as a legacy for future generations.     

Endemic predators, invasive prey and native diversity

Interactions between native diversity and invasive species can be more complex than is currently understood. Invasive ant species often substantially reduce diversity in the native ants diversity that act as natural control agents for pest insects. In Indonesia (on the island of Sulawesi), the third largest cacao producer worldwide, we show that a predatory endemic toad (Ingerophrynus celebensis) controls invasive ant (Anoplolepis gracilipes) abundance, and positively affects native ant diversity. We call this the invasive-naivety effect (an opposite of enemy release), whereby alien species may not harbour anti-predatory defences against a novel native predator. A positive effect of the toads on native ants may facilitate their predation on insect vectors of cacao diseases. Hence, toads may increase crop yield, but further research is needed on this aspect. Ironically, amphibians are globally the most threatened vertebrate class and are strongly impacted by the conversion of rainforest to cacao plantations in Sulawesi. It is, therefore, crucial to manage cacao plantations to maintain these endemic toads, as they may provide critical ecosystem services, such as invasion resistance and preservation of native insect diversity.     

Molecular characterization of Hepatozoon species in reptiles from the Seychelles

Hepatozoon parasites were examined for the first time in reptiles from the Seychelles Islands. Although both prevalence and intensity were low, Hepatozoon species were detected in individuals from 2 endemic species, the lizard Mabuya wrightii and the snake Lycognathophis seychellensis. This was confirmed using visual identification and through sequencing part of the 18s rRNA gene. Phylogenetic analysis indicates that the Hepatozoon on the Seychelles form a monophyletic lineage, although more data are clearly needed to stabilize estimates of relationships based on this marker.     

Life history evolution in Lodoicea maldivica (Arecaceae)

Lodoicea maldivica , a palm endemic to two small islands in the Seychelles group, has the largest seed in the plant kingdom. We present here an interpretation of the ecological and evolutionary significance of this seed in terms of the island environment where the species grows. We begin by reviewing the available information about the biology and ecology of Lodoicea and present some original data on the growth and development of Lodoicea in its native habitat. A remarkable feature of young plants is the enormous size of their leaves and the great length of their petioles, these being especially elongated when growing beneath the canopy. As a result, juvenile plants can reach a height of 15 m and hold their foliage in the forest canopy. This capacity to produce such an enormous juvenile plant is related in part to the large food reserves in the seed. We suggest that Lodoicea evolved from a more typical borassoid palm (perhaps a plant like Borassus aethiopum which is widespread in the savannas of Africa) and propose two hypotheses to explain why this occurred. According to the 'shade hypothesis', increasingly humid conditions on the Seychelles led to strong selection for plants with the tallest seedlings, since these would be the most likely to establish successfully under the low light conditions prevailing on the forest floor of closed forest. The 'sibling competition hypothesis' postulates that the island populations of the ancestral palm lost any means for seed dispersal, and their seeds simply fell to the foot of the parent tree. This resulted in a strong selection pressure for reduced fecundity. The two hypotheses are not mutually exclusive, and together they may help us to understand many of the unusual features of this remarkable species.     

Weak divergence among African, Malagasy and Seychellois hinged terrapins (Pelusios castanoides, P. subniger) and evidence for human-mediated oversea dispersal

Using phylogenetic and haplotype network analyses of 2036 bp of mitochondrial DNA, we compare samples of the two hinged terrapin species Pelusios castanoides and P. subniger from continental Africa, Madagascar and the Seychelles to infer their biogeography. Owing to the long independent history of Madagascar and the Seychelles, the populations from those islands should be deeply divergent from their African conspecifics. Seychellois populations of the two species are currently recognized as Critically Endangered endemic subspecies. However, even though we found within P. subniger evidence for a cryptic species from the Democratic Republic of the Congo, all other samples assigned to this species were undifferentiated. This suggests that Malagasy and Seychellois populations of P. subniger were introduced by humans and that the Seychellois subspecies P. s. parietalis is invalid. This has implications for current conservation strategies for the Critically Endangered Seychellois populations and suggests that measures should rather focus on endemic species. The situation of P. castanoides could be different. Samples from Madagascar and the Seychelles are weakly, but consistently, differentiated from continental African samples, and Malagasy and Seychellois samples are reciprocally monophyletic in maximum likelihood analyses. However, due to a lack of samples from central and northern Mozambique and Tanzania, we cannot exclude that identical continental haplotypes exist there.     

Strategic Management of an Invasive Ant‐scale Mutualism Enables Recovery of a Threatened Tropical Tree Species

Mutualisms between invasive ants and honeydew‐producing insects can have widespread negative effects on natural ecosystems. This is becoming an increasingly serious problem worldwide, causing certain ecosystems to change radically. Management of these abundant and influential mutualistic species is essential if the host ecosystem is to recover to its former non‐invaded status. This negative effect is particularly prevalent on some tropical islands, including Cousine Island, Seychelles. On this island, the invasive ant Pheidole megacephala has caused serious indirect damage to the threatened native Pisonia grandis trees via a mutualism with an invasive scale insect, Pulvinaria urbicola. We aimed to suppress the ant, thereby decoupling the mutualism and enabling recovery of the Pisonia trees. We treated all areas where ant pressure was high with a selective formicidal bait, which was deployed in custom‐made bait stations designed to avoid risk of treatment to endemic fauna. In the treated area, ant foraging activity was reduced by 93 percent and was followed by a 100 percent reduction in scale insect density. Abundance of endemic herbivorous insects and herbivorous activity increased significantly, however, after the decline in mutualistic species densities. Despite the native herbivore increase, there was considerable overall improvement in Pisonia shoot condition and an observed increase in foliage density. Our results demonstrate the benefit of strategic management of highly mutualistic alien species to the native Pisonia trees. It also supports the idea that area‐wide suppression is a feasible alternative to eradication for achieving positive conservation management at the level of the forest ecosystem.     

A phylogenetic analysis of the palm subtribe Oncospermatinae (Arecaceae) based on morphological characters

Subtribe Oncospermatinae (Arecaceae: Arecoideae: Areceae) is a diverse group of spiny Old World palms. The subtribe includesOncosperma, a widespread Asian genus of five species, along with seven monotypic genera, all endemic to the Seychelles and Mascarene Islands of the western Indian Ocean. A phylogenetic analysis was conducted in order to test the monophyly of subtribe Oncospermatinae with respect to other Old World genera of tribe Areceae. A matrix of 38 morphological characters was scored for 29 taxa, including 11 species of the Oncospermatinae. A single most parsimonious tree was found, resolving the subtribe as a polyphyletic group of two distinct clades. One clade containingAcanthophoenix, Deckenia, Oncosperma, andTectiphiala was placed as sister to a large group that includes members of subtribes Archontophoenicinae, Arecinae, Iguanurinae, and Ptychospermatinae. The other clade of Oncospermatinae, including the Seychelles endemic generaNephrosperma, Phoenicophorium, Roscheria, andVerschaffeltia, was resolved as sister to the Madagascar endemic subtribe Masoalinae, and may have arisen in the western Indian Ocean region.     

Response of a threatened giant millipede to forest restoration

Understanding the response of threatened and functionally important island invertebrate species to plant community restoration is essential for the successful conservation of these invertebrate species. The Seychelles giant millipede (SGM), Sechelleptus seychellarum, is a threatened and functionally important macro-detritivore endemic to the Seychelles granitic islands. Here, we studied the response of this species to the ongoing forest restoration programme on Cousine Island, Seychelles. This study was conducted over an 11 year period, representing the floral succession of a formerly degraded open scrubland area, dominated by alien plants, to a closed canopy forest, dominated by planted indigenous trees. While the time span of this study was insufficient for vegetation height in the restored area to equal that in the reference natural forest, canopy closure was nevertheless comparable. We found SGM density to be an order of magnitude lower in the restored site compared to the natural forest. In contrast, SGM physical condition improved significantly in the restored site, as vegetation structure increased. Furthermore, SGM behaviour in the restored site switched from a predominantly walking to a dominantly feeding behaviour over the study period, resulting in the forest restoration programme on Cousine increasing the foraging area of the SGM by 43 %. Competition for key resources, e.g. food and day-time refuges, are suggested as possible factors limiting SGM numbers in the restored forest.