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.     

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.     

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.     

Longevity and survival of the Endangered Seychelles Magpie Robin Copsychus sechellarum

The Seychelles Magpie Robin Copsychus sechellarum was once one of the most threatened birds in the world, but was downgraded from Critically Endangered to Endangered after a long-term recovery programme was success- fully implemented. Comprehensive long-term monitoring of this species was conducted on the islands of Cousin and Cousine over an 18-year period. We report here on the species longevity and annual survival at these two sites. The oldest recorded individual was a male who died on Cousine Island on 28 September 2000 at just under 16 years old. This individual was recorded to have hatched on Frégate Island on 3 January 1985, before being translocated to Cousine Island in 1995. Mean annual survival rates over an 18-year period were 81.6% on Cousin and 77.9% on Cousine islands. A decrease in annual survival was noted with increasing population size on both islands (Cousin: t = ?3.09, p < 0.05; Cousine: t = ?2.71, p < 0.05), which is a likely consequence of increased territory disputes and competition for food.     

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.     

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.     

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.     

First record of the fledging period of Temminck's Courser,Cursorius temminckii

The Seychelles Fody, Foudia sechellarum, is a ploceid weaver occurring naturally on three islands in the Seychelles group in the Indian Ocean. The population on Cousine Island was studied between 30 June and 25 August 1997. The size of the population on the island was estimated at 458–614 individuals and densities varied in different habitat types. As Seychelles Fodies in non-breeding plumage are difficult to sex, we provide sexing criteria based on wing length. Breeding pairs form small, probably temporary, territories that are defended by both partners against other fodies, including the introduced Madagascar Fody, Foudia madagascariensis. The Seychelles Fody often breeds semi-colonially and we observed up to five nests close together. The birds are socially monogamous and both sexes share in nest building, nest defense and provisioning of the young. Only females incubate. Many nests were deserted before eggs were laid, including some that had been accepted by the female. Additional males and females were sometimes seen helping provision the young, but this cooperative breeding behaviour appeared to be uncommon. Non-breeding individuals congregate in large flocks, sometimes joined by breeding birds.     

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.     

Predation risk affects trade-off between nest guarding and foraging in Seychelles warblers

The fitness costs of egg loss for Seychelles warblers (Acrocephalus sechellensis)on Cousin Island are considerable because warblers have a single-eggclutch and no time to lay a successful replacement clutch. Onthe islands of Cousin and Cousine, with equal densities of Seychellesfodies (Foudia sechellarum), nearly 75% of artificial eggs placedin artificial nests were predated by fodies after 3 days. OnAride Island with no fodies present, loss of artificial eggswas not observed. Female warblers incubate the clutch, and malewarblers guard the clutch when females are absent. Deterrenceof fodies by male warblers is efficient: loss rate of eggs fromunattended warbler nests was seven times as high as from attendednests, and the more nest guarding, the lower the egg loss andthe higher the hatching success. Egg loss is independent ofthe amount of incubation by females. There is no trade-off betweenincubating and foraging by females. Nest guarding competes withforaging by males, and this trade-off has a more pronounced effecton egg loss when food availability is low. The transfer of breeding pairsfrom Cousin to either Cousine with egg-predating fodies or toAride without fodies allowed us to experimentally investigatethe presumed trade-off between nest guarding and foraging. OnCousine, individual males spent the same amount of time nestguarding and foraging as on Cousin, and egg loss was similarand inversely related to time spent nest guarding as on Cousin.Males that guarded their clutch on Cousin did not guard theclutch on Aride but allocated significantly more time to foragingand gained better body condition. Loss of warbler eggs on Aridewas not observed. Time allocation to incubating and foragingby individual females before and after both translocations remainedthe same.     

Daughters on request: about helpers and egg sexes in the Seychelles warbler

The Seychelles warbler (Acrocephalus sechellensis) was an endangered endemic of the Seychelles islands where, until 1988, the entire population of ca. 320 birds was restricted to the one island of Cousin Island (29 ha). Additional breeding populations were successfully established on the islands of Aride (68 ha, 1988) and Cousine (26 ha, 1990) and now with the existence of ca. 2000 warblers on three islands the conservation status of the warbler has improved from endangered to vulnerable. Emigration from the island is extremely rare, so birds that disappeared were known to have died. Almost every bird on Cousin Island has been individually colour ringed and monitored throughout all breeding attempts during a 17-year period (1985-2002; total ca. 2400 birds). These birds were also blood sampled for molecular parentage and sex analyses. Therefore the lifetime reproductive success of many birds is known. Although warblers can breed independently in their first year, some individuals remain in their natal territory as subordinates, and often help by providing nourishment to non-descendant offspring. The frequency of 'helping' is affected by habitat saturation and variation in territory quality (insect prey availability). The long-term benefits of helping are higher for daughters than for sons, and it is therefore no wonder that most helpers are daughters from previous broods. Furthermore, on low-quality territories breeding pairs raising sons gain higher fitness benefits than by raising daughters, and vice versa on high-quality territories. Female breeders adaptively modify the sex of their single-egg clutches according to territory quality: male eggs on low quality and female eggs on high quality. The Seychelles warbler is a beautiful example of behavioural and life-history adaptations to restricted circumstances.     

Food and density limitations of the Seychelles Magpie Robin,Copsychus sechellarum,on Cousine Island

The critically endangered Seychelles Magpie Robin, Copsychus sechellarum, is one of the rarest birds in the world. At the end of December 1999, there were 88 individuals distributed on four small granite islands: Fregate, Aride, Cousin and Cousine. Little is known of Magpie Robin ecology in its natural habitat. Studies carried out on Aride, Cousin and Cousine are therefore valuable as each island is dominated by native woodland. The composition of Magpie Robin diet was compared between Fregate and Cousine. An exotic species of cockroach, Pycnoscelus indicus, and dropped fish were found to be considerably more important prey items on Cousine. Observations of chick food provisioning indicated habitat quality differences between the territories. Invertebrate and vertebrate food resources available to the Magpie Robin were sampled on Cousine between 1997 and 1998. Fourteen sites were sampled for invertebrates and fifty-two species from fifteen taxa were identified. There were significant differences between the mean number of species recorded per site and the mean number of animals found at each site. Skink density was estimated at between 1219-1516/ha and 354-538/ha for Mabuya sechellensis and Mabuya wrightii respectively. Invertebrate diversity and abundance was greatest in areas dominated by closed-canopy woodland on or near the coastal plain. These results explained the current distribution of Magpie Robin territories. The invertebrate abundance data were used to estimate the carrying capacity of the island for the Magpie Robin. Cousine could theoretically support up to six breeding pairs but it is questionable that a population of this size could be self-sustaining in the long-term.     

Recent invasion of European red foxes (Vulpes vulpes) on to Fraser Island (K'gari) and South Stradbroke Island

Invasive predators are globally significant drivers of threatened fauna population decline and extinction, and the early detection of new incursions is critical to the chances of successful predator eradication and fauna conservation. Here, we provide evidence of the recent invasion of European red foxes (Vulpes vulpes) on to two large and internationally significant islands off the southeast coast of Queensland, Australia – Fraser Island (K'gari) and South Stradbroke Island. From camera trap footage collected on Fraser Island since 2009, foxes have now been observed on seven different occasions between 2012 and 2016. Two scats collected on South Stradbroke Island in 2013 and 2014 tested positive for fox DNA (and negative for Canis spp. DNA), with fox presence confirmed by subsequent camera trap footage in 2016. These data confirm the recent incursion of foxes on to these islands and suggest that small populations now exist there. Fraser Island and South Stradbroke Island represent key RAMSAR wetland areas of refuge for populations of multiple threatened fauna that have never been previously been exposed to foxes. Fox impacts on these fauna can only be expected to increase without management intervention to eradicate them before they become widespread.     

Conservation genetics of critically endangered Crepidiastrum grandicollum (Asteraceae) and two closely related woody species of the Bonin Islands,Japan

Crepidiastrum grandicollum is a critically endangered insular endemic herb found only on two oceanic islands of the Bonin Islands in Japan, namely Chichijima Island and Anijima Island. It is explicitly threatened by herbivory pressure from introduced animals. In 2009, a conservation program for C. grandicollum was begun to ensure its future persistence. To provide further information for conservation planning, we investigated the genetic diversity of C. grandicollum using 13 novel microsatellite markers in 55 individuals from four wild populations and an ex situ living collection. Two closely related woody species were also included for interspecific comparison: seven individuals of Crepidiastrum ameristophyllum and 13 of Crepidiastrum linguifolium. The 13 markers were applicable to all three species and identified 129 alleles in total. We found a clear genetic differentiation between C. grandicollum from Anijima Island and Chichijima Island. Crepidiastrum grandicollum also had low expected heterozygosity and allelic richness in populations compared to the two closely related species. Reconstructed divergence history suggested that differentiation between the islands had occurred several thousand generations ago. We suggest separate conservation units for C. grandicollum on Anijima Island and Chichijima Island given the clear (and putatively historical) genetic differentiation, which may result in speciation in the future.

     

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.     

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.     

Colonization of Colliery Spoil Heaps by Millipedes (Diplopoda) and Terrestrial Isopods (Oniscidea) in the Sokolov Region, Czech Republic

Successive development of millipede and terrestrial isopod assemblages in colliery spoil heaps in the Sokolov region (northwest Bohemia, Czech Republic) was studied during the years 1993–1998. Younger colliery spoil heaps were characterized by a lower number and low density of millipede species. The millipedes Polydesmus inconstans, Polydesmus testaceus, and Craspedosoma rawlinsii represented the main pioneer species. Alder afforestation proved to be more suitable for the recolonization of millipede populations, especially for the predominant Julus scandinavius. Terrestrial isopods colonized all parts of the heaps, including the youngest open plots. The most frequent isopod, Trachelipus rathkii, is a colonizing species with an apparently high adaptability to different environmental conditions. The isolated character of the extensive heap areas, together with existing extreme soil conditions, resulted in slow recolonization and a successional development of stable populations of these animals. However, the relatively high densities of millipede and isopod assemblages in suitable alder afforestation can represent further potential sources for colonization of other open‐heap areas.     

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.     

A conceptual model of ecosystem restoration triage based on experiences from three remote oceanic islands

A conceptual model, that illustrates restoration, ecological landscaping, rehabilitation and regreening, is developed. It considers biocentric, historical, aesthetic and engineering aspects. The term ecosystem restoration triage is used because the first step is to decide whether to do nothing (because, on the one hand, the system is too degraded to warrant restoration, or, on the other, because biological integrity is relatively intact and therefore either none, or minimal, restoration is required) or to do something (because restoration is worthwhile, urgent and feasible). This approach hinges on the definition that restoration in the strictist sense is a biocentric activity that returns the original compositional, structural and functional diversity, along with its dynamics and natural evolutionary potential. Original is a difficult qualifier as it depends on just how far back in time we go. Where human values are involved, this is not restoration in the pure sense of restoring ecological integrity, but is ecological landscaping, rehabilitation or regreening. Experience from three remote oceanic islands [Easter Island, Cousine Island (Seychelles), Marion Island (Sub-Antarctic)] and which represent near extremes of this model are used to illustrate it.     

From small Maria to massive Campbell: Forty years of rat eradications from New Zealand islands

Abstract

Over the last four decades the eradication of rats from islands around New Zealand has moved from accidental eradication following the exploratory use of baits for rat control to carefully planned complex eradications of rats and cats (Felis catus) on large islands. Introduced rodents have now been eradicated from more than 90 islands. Of these successful campaigns, those on Breaksea Island, the Mercury Islands, Kapiti Island, and Tuhua Island are used here as case studies because they represent milestones for techniques used or results achieved. Successful methods used on islands range from bait stations and silos serviced on foot to aerial spread by helicopters using satellite navigation systems. The development of these methods has benefited from adaptive management. By applying lessons learned from previous operations the size, complexity, and cost effectiveness of the campaigns has gradually increased. The islands now permanently cleared of introduced rodents are being used for restoration of island‐seabird systems and recovery of threatened species such as large flightless invertebrates, lizards, tuatara, forest birds, and some species of plants. The most ambitious campaigns have been on remote subantarctic Campbell Island (11 300 ha) and warm temperate Raoul Island (2938 ha), aimed to provide long‐term benefits for endemic plant and animal species including land and seabirds. Other islands that could benefit from rat removal are close inshore and within the natural dispersal range of rats and stoats (Mustela erminea). Priorities for future development therefore include more effective methods for detecting rodent invasions, especially ship rats (Rattus rattus) and mice (Mus musculus), broader community involvement in invasion prevention, and improved understanding of reinvasion risk management.