Cousin Island Research Articles

Exploration is dependent on reproductive state, not social state, in a cooperatively breeding bird.

Personality is an intriguing phenomenon in populations because it constrains behavioral flexibility. One theory suggests that personality could be generated and maintained if dependent on asset protection. It is predicted that trade-offs with fitness expectations and survival probability encourage consistent behavioral differences among individuals (personality). Although not mutually exclusive, the social niche specialization hypothesis suggests that a group of individuals that repeatedly interact will develop personality to avoid costly social conflict. The point at which behavioral consistency originates in the social niche hypothesis is still unclear, with predictions for development after a change in social status. In the facultative cooperatively breeding Seychelles warbler (Acrocephalus sechellensis), residing on Cousin Island, breeding vacancies are limited and this forces individuals into different social roles. We used this system to test whether reproductive and social state predicted among-individual differences in exploration. We had 2 predictions. First, that an individual's start in life can predict personality, whereby young individuals with a good start to life (associated with early age reproduction and earlier onset survival senescence) are fast explorers, suggesting reproductive state-dependence. Second, that an individual's social status can predict personality, whereby dominant individuals will be fast explorers, suggesting that the behavior is social state-dependent. Neither of the behaviors was associated with social state and social state did not affect behavioral consistency. However, novel object exploration was associated with a proxy of reproductive state. Our results provide further support for state being a mechanism for generating individual differences in behavior.

No Association between Personality and Candidate Gene Polymorphisms in a Wild Bird Population.

Consistency of between-individual differences in behaviour or personality is a phenomenon in populations that can have ecological consequences and evolutionary potential. One way that behaviour can evolve is to have a genetic basis. Identifying the molecular genetic basis of personality could therefore provide insight into how and why such variation is maintained, particularly in natural populations. Previously identified candidate genes for personality in birds include the dopamine receptor D4 (DRD4), and serotonin transporter (SERT). Studies of wild bird populations have shown that exploratory and bold behaviours are associated with polymorphisms in both DRD4 and SERT. Here we tested for polymorphisms in DRD4 and SERT in the Seychelles warbler (Acrocephalus sechellensis) population on Cousin Island, Seychelles, and then investigated correlations between personality and polymorphisms in these genes. We found no genetic variation in DRD4, but identified four polymorphisms in SERT that clustered into five haplotypes. There was no correlation between bold or exploratory behaviours and SERT polymorphisms/haplotypes. The null result was not due to lack of power, and indicates that there was no association between these behaviours and variation in the candidate genes tested in this population. These null findings provide important data to facilitate representative future meta-analyses on candidate personality genes.

Reef fishes recruited at midwater coral nurseries consume biofouling and reduce cleaning time in Seychelles, Indian Ocean

In coral reef restoration, coral gardening involves rearing coral fragments in underwater nurseries prior to transplantation. These nurseries become fish-aggregating devices and attract biofouling. We hypothesised that: (1) the presence of corals at a nursery is critical to recruit fish assemblages and (2) the recruited fish assemblages control biofouling, reducing person-hours invested in nursery cleaning. Three midwater coral nurseries were deployed at 8 m depth for 27 months within the marine protected area of Cousin Island Special Reserve, Seychelles, Indian Ocean. Each nursery consisted of a 6 m×6 m PVC pipe frame, layered with a recycled 5.5-cm-mesh tuna net. Human cleaning effort was calculated based on daily dive logs. Nursery-associated fish assemblages and behaviour were video-recorded prior to harvesting corals after a 20-month growth period and seven months post-coral harvesting. The density (ind. m–2) of blue-yellow damselfish Pomacentrus caeruleus was 12–16 times higher when corals were present than when corals were absent at the nurseries. Fish assemblages recruited into the nurseries included three trophic levels, from herbivores to omnivores, in six families: Ephippidae, Pomacentridae, Labridae (Scarinae), Gobiidae, Siganidae and Monacanthidae. Higher abundance of large fish (total number of individuals) resulted in 2.75 times less person-hours spent in nursery cleaning. These results have important implications for cost-effective coral reef restoration.

Spatial variability in habitat structure and heterogenic coral reef fish assemblages inside a small-scale marine reserve after a coral mass mortality event

Coral reefs at the inner granitic islands in the Seychelles were heavily affected by the worldwide bleaching event in 1998, which led to subsequent coral mortality and widespread phase shifts to macroalgae dominated reefs. In this study, five sites within a small, but well enforced marine reserve at Cousin Island, were investigate using various methods to explore differences in coral habitat quality, coral recruitment, fish assemblages, key invertebrate grazers, and rugosity. The objective of the study was to collect a broad set of scientific data, which could be useful to describe linkage between coral reef and fish assemblages after a large-scale disturbance, as well as for future management decisions regarding marine resources, in terms of MPA protection and recovery abilities. The results showed high spatial variation in coral coverage between sites (from 1.5% to 43.2%), which were higher than previously reported, as well as high variation in dispersal of coral recruits. Furthermore, there were large heterogenic differences in fish densities and composition, which were directly linked to coral habitat quality, e.g. total fish abundance was 15 times higher on sites with high coral coverage in comparison to sites with low coral cover. In summary, this study demonstrates that coral reef habitat and fish assemblage may display high spatial variability and heterogenic differences after large-scale disturbances and suggests that potential recovery from coral mass mortality may occur in a non-linear and patchy procedure, which in turn may depend on underlying stocastical processes that affect coral recruitment and survivorship.

Saving a tropical ecosystem from a destructive ant-scale (Pheidole megacephala, Pulvinaria urbicola) mutualism with support from a diverse natural enemy assemblage

Ants can disrupt the natural biological control of serious hemipteran pests by interfering with natural enemies, resulting in a change in ecosystem functioning. We focus here on interference by a highly invasive ant Pheidole megacephala on the regulation of a tree destroying hemipteran scale insect Pulvinaria urbicola on Cousine Island in the Seychelles archipelago, a tropical island ecosystem. We show how a diverse natural enemy assemblage contributes substantially to the collapse of the ant-scale mutualism following managed ant suppression. Natural enemy abundance and species richness increased significantly after ant suppression, with varying responses among the different functional guilds. Primary parasitoids coexisted with tending ants before ant suppression, but could not regulate the enormously high scale densities alone. After ant suppression, a significant increase in predators caused a collapse of the scale population. Guilds external to the mutualism were also affected, with primary parasitoids of various non-hemipteran taxa also increasing, which contributed significantly to the recovery of the community to its pre-invasion composition. Our results highlight the far-reaching and pervasive effects of the hemipteran-tending invasive ant within the natural enemy assemblage. In turn, we also illustrate the potential to restore the tropical ecosystem by encouraging an array of natural enemies through precision management of the ant.

Population dynamics of a threatened giant millipede: implications for restoration

AbstractThe population dynamics of threatened invertebrates have important implications for their conservation and restoration. 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 population dynamics of the SGM from 1998 to 2009 on Cousine Island, Seychelles, to make practical restoration recommendations. Large fluctuations in millipede densities were found between 1998 and 2009. In 2002, 2003, 2005, and 2007 millipede densities were low, while densities were high in 1998 and 2009. Although the SGM is active all year round, millipede surface activity was positively correlated with rainfall, with millipede density high during the wet NW monsoon period (i.e., October to April) and low during the SE trade wind period (i.e., May to September). Female:male:juvenile ratios were ∼3:1:1. The implications of these results for restoration are that translocations should preferably be done in years of high millipede densities and during the wet season. Furthermore, chemical control of the invasive ant Pheidole megacephala, which is currently being carried out on Cousine Island and in future could be conducted on other Seychelles islands, should preferably be done during the low rainfall months, as the SGM readily consumes the hydramethylnon‐based bait.

Beyond vegetation-based habitat restoration for a threatened giant Spirostreptid millipede

Vegetation-based restoration efforts often fail to provide suitable habitat for many invertebrates. Restoring habitat for target invertebrates requires an understanding of their resource and condition requirements. The Seychelles giant millipede (SGM), Sechelleptus seychellarum, is a functionally important millipede which is Red Listed as ‘Vulnerable’. Here, we studied the habitat requirements of this macro-detritivore in relation to the ongoing forest restoration programme on Cousine Island, Seychelles. SGM spatial density did not significantly correlate with edaphic and litter properties, but did positively correlate with the toposcape (i.e. elevation and granitic rock cover). Granite rock crevices in forest covered areas were important diurnal refuges for the SGM, as microclimate conditions in non-forest covered rock refuges were unsuitable. SGM physical condition was significantly lower in non-shaded crevices compared to those shaded by forest. Low granite rock cover in the restored forest limited the SGM colonisation of this area in large numbers, despite canopy cover in the restored forest being comparable with that in the reference natural forest. As most restoration practices are primarily vegetation-based, this study demonstrates that such an approach can be inadequate for restoring habitat for target invertebrates, as many species habitat requirements extend beyond that of vegetation. For the SGM, selecting restoration sites that already have abundant rock cover would be the most practical way to increase SGM habitat through forest restoration practices. Taking into consideration the habitat requirements of target invertebrates can help in setting or redirecting restoration goals and thus enhance the conservation value of such practices.

A Behavioural Ecology Approach to Assessing the Effect of Alien Vegetation on a Threatened Giant Millipede

The importance of taking into account behavioural ecology when assessing the impacts of habitat deterioration on threatened species is evaluated. The Seychelles giant millipede (SGM), Sechelleptus seychellarum, is IUCN Red-Listed as “Vulnerable”. Here, we studied the effect of alien bamboo and coconut on the behavioural ecology of this species on Cousine Island, Seychelles. Coconut did not affect SGM density, but negatively affected its foraging behaviour, whereas bamboo negatively affected both its density and foraging behaviour. The SGM showed feeding preferences for Pisonia grandis and Ficus sp. leaf litter types. Alien bamboo and coconut pose a varied threat to the SGM, and their removal and replacement by indigenous forest species (e.g. P. grandis and Ficus sp.) should form part of an island’s restoration programme.

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.

The Art and Science of Restoration

The Art and Science of Restoration

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

AbstractMutualisms 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.

Impact of a mutualism between an invasive ant and honeydew-producing insects on a functionally important tree on a tropical island

Mutualisms between invasive ants and honeydew-producing Hemiptera have the potential to result in unusually high population levels of both partners, with subsequent major changes to ecosystem composition and dynamics. We assessed the relationship between the invasive ant, Pheidole megacephala, and its hemipteran mutualists, Dysmicoccus sp. and Pulvinaria urbicola, on Cousine Island, Seychelles. We also assessed the impacts of the mutualism on the condition of the hemipteran host plant, Pisonia grandis, a native and functionally important tree species. There was a strong positive relationship between Ph. megacephala activity and hemipteran abundance, and the exclusion of ants from Pi. grandis resulted in a significant decline in Pu. urbicola abundance. High abundance of the mutualists was strongly associated with damage to the Pi. grandis forest. This indicates that the mutualism is contributing to the massive increase in the population levels of the mutualist species, and is intensifying their impacts on the island. The widespread trophobiosis and its associated high densities of mutualists pose serious threats to the ecosystem, highlighting the need to control the ant and associated hemipteran populations.

Tropical island recovery: Cousine Island, Seychelles

Tropical island recovery: Cousine Island, Seychelles

Hawksbill turtle monitoring in Cousin Island Special Reserve, Seychelles: an eight-fold increase in annual nesting numbers

Results of hawksbill turtle Eretmochelys imbricata nest monitoring on Cousin Island, Seychelles, indicate an 8-fold increase in abundance of nesting females since the early 1970s when the population was highly depleted. From 1999 to 2009, the population increased at an average rate of 16.5 turtles per season. Females were individually tagged, and nesting data were derived from indirect evidence of nesting attempts (i.e. tracks) and actual turtle sightings (56 to 60% of all encounters). Survey effort varied over the years for a variety of reasons, but the underlying trends over time are considered robust. To overcome biases associated with variable survey effort, we estimated population changes by fitting a Poisson distribution to data on numbers of times each individual was seen at this breeding site in a season. This was used to estimate unseen individuals, and hence the total number of nesting females each season. The maximum number of individuals emerging onto Cousin Island to nest within a single season was estimated to be 256 (2007 to 2008) compared to 23 in 1973. Tag returns indicate that many turtles nest on both Cousin and Cousine Islands (2 km apart), and that some inter-island nesting also occurs between Cousin and more remote islands within the Seychelles.

Population size and home range of black-naped hares Lepus nigricollis nigricollis on Cousin Island (Seychelles, Indian Ocean)

Article Population size and home range of black-naped hares Lepus nigricollis nigricollis on Cousin Island (Seychelles, Indian Ocean) was published on January 1, 1994 in the journal Mammalia (volume 58, issue 4).

Nesting success of White Terns and White-tailed Tropicbirds on Cousine Island, Seychelles

This study investigates the breeding success of two tropical seabirds that exploit dissimilar nesting habitats on Cousine Island in the Seychelles archipelago, the White Tern Gygis alba and the White-tailed Tropicbird Phaethon lepturus, which nest in trees and in crevices on the ground, respectively. Both species have a clutch of one egg. Over a 23-month study period, the outcomes of 134 nesting attempts by White Terns and 285 by White-tailed Tropicbirds were followed. White Terns produced an average of 0.4 chicks per attempt, significantly more than that of White-tailed Tropicbirds (0.25). Hatching success did not differ between the two species, but fledging success of White Terns (62%) was significantly higher than that of White-tailed Tropicbirds (43%). Nesting success of White-tailed Tropicbirds may be less successful than White Terns because their ground nests are at risk to purely terrestrial predators, in addition to predators that are both terrestrial and arboreal. Many nests of White-tailed Tropicbirds failed during the first two weeks of the incubation and nestling periods, but the reasons for this are unclear. Food availability may influence the reproductive success of both species.

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.

GRANDPARENT HELPERS: THE ADAPTIVE SIGNIFICANCE OF OLDER, POSTDOMINANT HELPERS IN THE SEYCHELLES WARBLER

The possibility that older, often nonreproductive, individuals may engage in kin-directed cooperative behavior has been largely overlooked in the study of cooperative breeding. Here, we describe and investigate the adaptive significance of such "grandparent" helpers in the Seychelles warbler, the first bird species in which this phenomenon has been observed. On Cousin Island, over a period of 24 years, a significant proportion (13.7%) of females, but few males (3.0%), was deposed from dominant positions. Deposed females were replaced by related females. However there was no evidence that older, senescent females were stepping aside to gain greater fitness benefits by increasing the reproductive success of their offspring, rather than breeding themselves; deposed females were not postreproductive, nor was being deposed linked to age or reproductive senescence. Of the deposed females, 68% became subordinates and helped to raise group offspring, accounting for ca. 10% of subordinates in any year. Demoted females were related (r= 0.24) to the group offspring and, consequently, could gain indirect benefits through helping. As direct benefits appeared to be limited, we suggest that indirect benefits have driven the evolution of such "grandparent helpers." This study now provides evidence for a new route to cooperative breeding in birds.

The role of exotic plants in the invasion of Seychelles by the polyphagous insect Aleurodicus dispersus: 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 Mahe, 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 Mahe, 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.

Phase shifts and the role of herbivory in the resilience of coral reefs

Cousin Island marine reserve (Seychelles) has been an effectively protected no-take marine protected area (MPA) since 1968 and was shown in 1994 to support a healthy herbivorous fish assemblage. In 1998 Cousin Island reefs suffered extensive coral mortality following a coral bleaching event, and a phase shift from coral to algal dominance ensued. By 2005 mean coral cover was <1%, structural complexity had fallen and there had been a substantial increase in macroalgal cover, up to 40% in some areas. No clear trends were apparent in the overall numerical abundance and biomass of herbivorous fishes between 1994 and 2005, although smaller individuals became relatively scarce, most likely due to the loss of reef structure. Analysis of the feeding habits of six abundant and representative herbivorous fish species around Cousin Island in 2006 demonstrated that epilithic algae were the preferred food resource of all species and that macroalgae were avoided. Given the current dominance of macroalgae and the apparent absence of macroalgal consumers, it is suggested that the increasing abundance of macroalgae is reducing the probability of the system reverting to a coral dominated state.