Field collections reveal that São Tomé is the Afrotropical island with the highest diversity of drosophilid flies (Diptera: Drosophilidae)

Summary

The Drosophilid fauna has been less investigated in the Atlantic Afrotropical islands than in the Indian Ocean. Located about 250 km from the continent, the volcanic island of São Tomé has been colonized mostly by natural means, probably by the wind, since the emergence of the island about 15 million years ago, and presumably also by anthropogenic transportation of invasive and domestic species. To date, 37 different Drosophilid species have been mentioned from São Tomé. The present work extends this list to 80 species. The genera Zygothrica, Phorticella and Hypselothyrea are newly recorded from the island. Among these 80 species, only 12 are putatively introduced by human activities, suggesting the preponderance of natural arrivals. Compared to other islands, São Tomé harbours a high diversity of drosophilids. At least 14 species are supposed to be endemic. Future molecular comparisons between the island flies and their continental relatives will probably help to identify other endemic species. The high diversity observed in São Tomé is certainly due to the large size of the island, and to the presence of vast natural altitudinal forests offering a variety of possible habitats. Further collections are likely to lead to an increase of the species list. From now, São Tomé island appears as an excellent laboratory for studying the ecology and evolution of the Drosophila model.     

Seasonal and inter‐island variation in the foraging strategy of the critically endangered Red‐headed Wood Pigeon Columba janthina nitens in disturbed island habitats derived from high‐throughput sequencing

We studied the feeding ecology of the critically endangered Red‐headed Wood Pigeon Columba janthina nitens, a subspecies endemic to a very remote and highly disturbed oceanic island chain, the Ogasawara Islands. An analysis based on high‐throughput sequencing (HTS) was undertaken on 627 faecal samples collected over 2 years from two island habitats, and food availability and the nutrient composition of the major fruits were also estimated. The HTS diet analysis detected 122 food plant taxa and showed clear seasonal and inter‐island variation in the diet of the Pigeons. The results indicated a preference for lipid‐rich fruits, but the diet changed according to the availability of food resources, perhaps reflecting the foraging strategy of the Pigeons in isolated island habitats with poor food resources. Pigeons also frequently consumed introduced plants at certain times of year, perhaps compensating for the lack of preferred native food resources. However, the degree of dependence on introduced plants appeared to differ between the two island habitats, so the different impacts of introduced plant eradication on the foraging conditions for the Pigeons on each island should be considered. HTS diet analysis combined with field data may be useful for monitoring the foraging conditions of endangered species and may also inform an appropriate conservation strategy in oceanic island ecosystems with complicated food webs that include both native and introduced species.     

Habitat use and dispersal characteristic by <Emphasis Type="Italic">Stethophyma grossum</Emphasis>: the role of habitat isolation and stable habitat conditions towards low dispersal

Species in a highly fragmented environment, such as the intensively used agricultural landscapes of Europe, are expected to be in danger of extinction. We hypothesize according to Kisdi’s theory (Am Nat 159:579–596, 2002) that species in fragmented landscapes with isolated habitats in general tend to possess low dispersal. In order to verify this hypothesis we studied the movement patterns of Stethophyma grossum, a hygrophilous species of wetlands, by mark–release–recapture techniques in a landscape with scattered suitable habitats over 3 years. The study focused on the major population in this landscape (site #1) as dispersal behaviour was assumed to be greatest. Actually, marked individuals of S. grossum were never found in any further suitable habitats in close vicinity to site #1. Despite that the peatland meadow of study site #1 was all over covered with homogenous vegetation only 6% (1.8 ha) of the whole area (30 ha) were occupied by S. grossum. The mean recapture rate over 3 years amounted to 39% with no significant differences between males and females. Both covered little distances within their mean range size of 1.8 ha; the median distances were 36.91 m for males and 26.65 m for females. We confirm the hypothesis that sub-populations of species in longstanding naturally isolated habitats, which habitat conditions have been stable; evolved low dispersal with little movements which are routine movements to find mating partners or food.     

Current distribution and potential extent of the most invasive alien plant species on La Réunion (Indian Ocean,Mascarene islands)

Abstract La Réunion Island has the largest area of intact vegetation of the islands in the Mascarene archipelago. Biological invasions are the primary threat to biodiversity in the intact habitats of the island (those not already transformed by agriculture and urbanization). Our study aimed to identify areas to prioritize in managing invasive alien plants for biodiversity conservation. We used extensive surveys of 238 distinct untransformed areas on La Réunion to define the current distribution patterns of all invasive species. Using expert knowledge, we compiled maps of the current distribution of the 46 most widespread/important invasive plants at the habitat scale (identified according to vegetation structure). Data from 440 botanical relevés for the 20 most threatening invasive alien plant species across the island and climatic envelope models were used to derive climatic suitability surfaces; these were used to map potential distributions for these species. More than 10 species invade 16.7% of the remaining habitat. Five habitat types are invaded by 25 or more species, and eight have fewer than 10 invasive alien plant species. Cluster analysis based on presence/absence of species in the 18 habitat types produced eight groups of species that invade particular habitats. Potential distribution models show that some species have invaded large parts of their potential range (e.g. Fuchsia magellanica, Furcraea foetida, Hiptage benghalensis), whereas others have the potential to increase their range substantially (e.g. Clidemia hirta, Strobilanthes hamiltonianus, Ulex europaeus). Management implications are identified for both groups. Three broad groups of habitats were identified: (i) intact habitats with a low level of invasion (e.g. subalpine shrubland); (ii) moderately invaded habitats with varying levels of intactness (ranging from windward submountain rainforest to the Acacia heterophylla forest); and (iii) habitats with little remaining intact area and high levels of invasion (e.g. lowland rainforest). Different management interventions are appropriate for these three groups.     

Relationships between intra‐specific variation in seed size and recruitment in four species in two contrasting habitats

Large seeds contain more stored resources, and seedlings germinating from large seeds generally cope better with environmental stresses such as shading, competition and thick litter layers, than seedlings germinating from small seeds. A pattern with small‐seeded species being associated with open habitats and large‐seeded species being associated with closed (shaded) habitats has been suggested and supported by comparative studies. However, few studies have assessed the intra‐specific relationship between seed size and recruitment, comparing plant communities differing in canopy cover. Here, seeds from four plant species commonly occurring in ecotones between open and closed habitats (Convallaria majalis, Frangula alnus, Prunus padus and Prunus spinosa) were weighed and sown individually (3200 seeds per species) in open and closed‐canopy sites, and seedling emergence and survival recorded over 3 years. Our results show a generally positive, albeit weak, relationship between seed size and recruitment. In only one of the species, C. majalis, was there an association between closed canopy habitat and a positive seed size effect on recruitment. We conclude that there is a weak selection gradient favouring larger seeds, but that this selection gradient is not clearly related to habitat.     

Dispersal and microsite limitation of a rare alpine plant

Knowledge on the limitation of plant species’ distributions is important for preserving alpine biodiversity, particularly when the loss of alpine habitats because of global warming or land use changes is faster than colonization of new habitats. We investigated the potential of the rare alpine plant Campanula thyrsoides L. to colonize grassland sites of different suitability on a small mountain plateau in the Swiss Alps. A total of 15 experimental sites were selected according to their differences in habitat suitability for adult C. thyrsoides, which was measured by the Beals index. At each site we applied a disturbance treatment, added seeds at different densities and monitored the survival of seedlings over two consecutive years. The number of surviving seedlings was not positively related to habitat suitability for adult C. thyrsoides. Furthermore, C. thyrsoides appears to be strongly dispersal limited at the regional scale because seed addition to unoccupied habitats resulted in successful germination and survival of seedlings. Since an increase of seed density in already occupied sites did not affect the number of seedlings, we suggest that C. thyrsoides is microsite limited at the local scale. Microsite limitation is supported by the result that seedling survival of the species was enhanced in vegetation gaps created by disturbance. We conclude that C. thyrsoides may become endangered in the future if environmental changes cause local extinction of populations. An appropriate management, such as a disturbance regime for enhancing recruitment in existing populations, may ensure the long-term survival of this rare alpine plant species.     

Effects of browsing in relation to vegetation cover on common yew (<Emphasis Type="Italic">Taxus baccata</Emphasis> L.) recruitment in Mediterranean environments

Common yew (Taxus baccata L.) stands are recognized as prioritary habitats for biodiversity conservation within the European Union. The effects of browsing on the regeneration capacity and spatial dispersal of T. baccata recruits at the European southern limit of the species in the Mediterranean Basin have been herein studied. The efficacy of T. baccata recruitment has been evaluated at six localities in the Northern Sardinia mountains, which have similar altitude, climate, soil, and vegetation but have different types of uses (three were grazed by livestock and three were not). At each site, five habitats have been identified for T. baccata seed dispersal: reproductive female T. baccata canopy, reproductive female Ilex aquifolium canopy, non-fleshy-fruited tree canopy, fleshy-fruited shrubs, and open areas. The density of seedlings was found to be greater under fleshy-fruited trees (reproductive female T. baccata and I. aquifolium) than under shrubs, whereas the sapling density was higher in shrubby habitats, especially at grazed sites due to the mechanical protection afforded by the spiny shrubs against herbivores. Land use (LU) has been found to be the most important factor in determining the spatial distribution of seedlings and saplings in relation to forest habitats. Although browsers had an ephemeral but positive effect on seed germination through their trampling and the resultant scarification, this process eventually became ineffective as was shown by the occurrence of the lowest density of saplings in those habitats where the density of seedlings was the highest. The ultimate and most important effect of browsing was the sharp decrease in the density of saplings, and their almost complete extinction, in non-shrubby habitats. This study highlights the result that, in Mediterranean ecosystems, browsing constitutes the main negative factor on T. baccata seedling-sapling transition and furthermore confirms the necessity to preserve shrubby patches in the vicinity of reproductive female T. baccata and I. aquifolium to permit the regeneration of T. baccata in the presence of livestock. Moreover, at ungrazed sites, T. baccata is able to colonize non-shrubby shady habitats. The application of different management strategies to ungrazed and grazed sites should therefore be the main direction in the management and preservation of T. baccata stands in the Mediterranean region.     

Archipelago-wide episodic recruitment of the file fish <Emphasis Type="Italic">Pervagor spilosoma </Emphasis> in the Hawaiian Islands as revealed in long-term records

The endemic filefish, Pervagor spilosoma, has long been known to recruit infrequently but in large numbers to shallow water habitats in Hawaii. When it does recruit, as it did in 1944, 1975 and 1982–1987, it becomes the most abundant or one of the most abundant species on Hawaiian reefs. At these times of high abundance, juvenile individuals also become abundant in the off-shore pelagic environment and in deeper benthic habitats. The results of two previously published studies and this one show that the last heavy recruitment occurred throughout the Hawaiian Island chain in the period from 1982–1988. The earliest strong recruitment in this interval occurred on the leeward, or northwest Hawaiian Islands in 1982, then on Oahu in 1983, then on the island of Hawaii, at the southeast end of the island chain in 1985. At the Kaneohe Bay, Oahu, sampling site, strong recruitment occurred in 1983–1987. Fish that had recruited in the summer of a year survived less than a year at these shallow (<15 m) sites or moved off the reefs. No other fish species among the 30 species censused in a 20 year period at Kaneohe Bay has shown the extremes of abundance displayed by this species, nor did any other species recruit so strongly in the 1983–1987 period. The latter event suggests that the pattern of recruitment in this species was not solely due to a particular pattern of ocean circulation, because if that were the case, other species would also show strong recruitment. The rarity of large recruitment events in this species, the high density of the recruits, the length of individuals at recruitment, the observation by others of juveniles and possibly young adults in the water column, and the sequential nature of the recruitment down the length of the island chain may in part be the result of metamorphosis in the pelagic environment and a pelagic juvenile stage in this species.     

Assessing the role of off‐take and source–sink dynamics in the extinction of the amphidromous New Zealand grayling (Prototroctes oxyrhynchus)

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

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

Genetic variation of pantropical Terminalia catappa plants with sea‐drifted seeds in the Bonin Islands: suggestions for transplantation guidelines

The Bonin Islands are endowed with endemic species. However, these species are at risk of extinction because of the exuberance of invasive alien plants. Therefore, native plant species should be revegetated after eradicating alien plants. We investigated the genetic variation of Terminalia catappa populations in the Bonin Islands by using nuclear (n) microsatellites (simple sequence repeats [SSRs]) and chloroplast (cp) DNA. No significant differences were observed in the genetic diversity of nSSRs among 22 populations. However, recent bottlenecks were detected in three populations on the Chichijima Island group. nSSR variation and cpDNA haplotypes suggested the presence of two genetically distinct groups in the Mukojima and Chichijima Island groups and the Hahajima Island group. A similar genetic structure was observed in plants and animals in the Bonin Islands. Populations on the three islands, which were separated from other islands in each island group when the water depth was 50‐m lower than the present level, were dominated by unique nSSRs clusters, suggesting that historical changes in island connections during the Pleistocene era affected genetic substructuring. These results suggested that different factors contributed to the genetic structure of T. catappa on different geographic scales. At the whole‐island level, the genetic structure was determined by long‐distance seed dispersal by ocean currents. At the island‐group level, the genetic structure was determined by historical changes in island connections caused by changes in the sea level due to glacial–interglacial transition. These findings would help in establishing transplantation zone borders for revegetating T. catappa on the Bonin Islands.     

Recruitment in invertebrates with short-lived larvae: the case of the bryozoan <Emphasis Type="Italic">Disporella hispida</Emphasis> (Fleming)

Temporal and spatial recruitment patterns of the cyclostome bryozoan Disporella hispida were monitored using settlement plates arranged along three benthic communities of an artificial reef at Blanes (NE Spain, NW Mediterranean). At the study site, the species mainly inhabits semi-obscure caves. By studying recruitment over one year I first inferred the larval release period for D. hispida and described its temporal occurrence in the communities and stations studied. Secondly, I attempted to determine whether the predicted restricted dispersal may account for the species' distribution at the study site. To this purpose, I compared the distribution of early recruits (15 days old) with that of adults. I also investigated environmental factors which may affect the extent of larval dispersal, and described the effects of post-recruitment processes occurring over a 4-month period. The brooding period, inferred from the study of early recruitment, was linked to spring-increments and autumn-decrements of water temperatures. Early recruits were distributed non-randomly in the communities and stations studied, being most abundant in the habitats where adults live. Strong hydrodynamic events seemed to modify this pattern, allowing recruitment out of the parental communities, and may hinder settlement. Post-recruitment mortality events were likely to prevent colonisation of habitats where the species do not live. Overall, philopatry and low post-recruitment mortality in the parental communities appeared to be the main mechanisms determining the distribution of D. hispida at the study site. Electronic Publication     

Habitat use and potential interactions between the house mouse and lesser white-toothed shrew on an island undergoing habitat restoration

To forecast the potential impact of plant community and dry-stone wall restoration on an insular population of the lesser white-toothed shrewCrocidura suaveolens Pallas, 1811, shrew and house mouseMus musculus Linnaeus, 1758 abundances were assessed in 3 anthropogenic habitats on Béniguet Island, Brittany, France, by a standardised annual trapping system checked yearly for 9 years and in 6 “natural” habitats by trap-lines. The standardised trapping system showed that abundances of both species fluctuated synchronously for nine years, suggesting that interactions between the species had little impact if any on abundances. Mice were trapped in all habitats, but shrews only in “stone” habitats except for rare occurrences in one damp depression. Ruderal habitat was rarely used by either species. Data suggest that on Béniguet Island: (1)M. musculus is associated with anthropogenic habitats but is not as strictly tied to them as at nearby continental sites; (2)C. suaveolens is synanthropic, as has been reported in continental northern France; (3) shingle beaches and seashore food resources are particularly important forC. suaveolens; (4) the relationship betweenC. suaveolens andM. musculus could not be determined by the experiments and, if it exists at all, appears to be more competitive than predatory. Grassland restoration is unlikely to affect shrew populations. Dry-stone wall restoration may temporarily affect shrews but should ultimately benefit them.     

Using population viability analysis to predict the effect of fire on the extinction risk of an endangered shrub <Emphasis Type="Italic">Verticordia fimbrilepis</Emphasis> subsp. <Emphasis Type="Italic">fimbrilepis</Emphasis> in a fragmented landscape

The fragmentation of mediterranean climate landscapes where fire is an important landscape process may lead to unsuitable fire regimes for many species, particularly rare species that occur as small isolated populations. We investigate the influence of fire interval on the persistence of population fragments of the endangered shrub Verticordia fimbrilepis Turcz. subsp. fimbrilepis in mediterranean climate south-west of Western Australia. We studied the population biology of the species over 5 years. While the species does recruit sporadically without fire this occurs only in years with above average rainfall, so fire seems to be the main environmental factor producing extensive recruitment. Transition matrix models were constructed to describe the shrub’s population dynamics. As the species is killed by fire and relies on a seed bank stimulated to germinate by smoke, stochastic simulations to compare different fire frequencies on population viability were completed. Extinction risk increased with increasing average fire interval. Initial population size was also important, with the lowest extinction risk in the largest population. For populations in small reserves where fire is generally excluded, inevitable plant senescence will lead to local extirpation unless fires of suitable frequency can be used to stimulate regeneration. While a suitable fire regime reduces extinction risk small populations are still prone to extinction due to stochastic influences, and this will be exacerbated by a projected drying climate increasing rates of adult mortality and also seedling mortality in the post-fire environment.     

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

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

Global diversity of polychaetes (Polychaeta; Annelida) in freshwater

A literature review of Polychaeta (Annelida) including Aphanoneura (the oligochaete-like Aeolosomatidae and Potamodrilidae), living in freshwater yielded 168 species, 70 genera and 24 families representing all of the major polychaete clades, but less than 2% of all species. The best-represented families were, in order, Nereididae, Aeolosomatidae, Sabellidae, Spionidae and Histriobdellidae. Fourteen families were represented by a single species and genus. Regions supporting the highest diversity of freshwater polychaetes were in order, Palaearctic, Neotropical, Oriental, Nearctic, Australasian, and Afrotropical. More than half of all species and genera inhabitat lakes and rivers, followed by lagoons/estuaries, which have a high proportion of euryhaline species, and inland seas. Less common, atypical polychaete habitats include subterranean waters, the hyporheic zone of rivers and plant container habitats (phytotelmata). At least three distinct ecological/historical processes appear to account for the colonisation of continental waters: invasion of a clade prior to the break-up of Gondwana, as in Aphanoneura, Namanereis, Stratiodrilus, and Caobangia; relatively recent stranding of individual species (relicts); and the temporary visitation of euryhaline species. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Dynamics of colonies of the speckled ground squirrel (<Emphasis Type="Italic">Spermophilus suslicus</Emphasis> Guld., 1770) on the northern boundary of the habitat

Results of studies in 1999–2009 of the dynamics of five isolated colonies of the speckled ground squirrel (Spermophilus suslicus) at the northern boundary of the habitat of this species (Zaraiskii area, Moscow oblast) are given. An abrupt decrease in the number of this species in colonies was established, as was a multifold decrease (up to 70–90%) in suitable habitats due to the plowing of meadows and build-up of the area. The fragmentation of the optimum habitat of ground squirrels reached a stage where colonies can no longer exist as a stable autonomous system. The process of their extinction unfolds quickly: the number of animals in some colonies decreased five-six times over the last ten years. To preserve this species, it is necessary to create areas of steppefied meadows with regulated agricultural use.     

Postimpoundment Changes in Water Quality and Fish Stocks in Two Large West African Reservoirs (Manantali and Sélingué, Mali)

The recently established large reservoirs Manantali on the river Bafing, tributary to the Senegal River and Sélingué, located on the river Sankarani, tributary to the Niger River, have followed similar changes in limnology and fish populations as those established in West Africa 20 to 30 years ago. Basic water parameters, such as oxygen content, pH, electrical conductivity and temperature show seasonal variations largely determined by the changes in wet and dry seasons. Both waterbodies are monomictic, with water mixing taking place in January, the coldest period of the year. Pre-impoundment forest clearence of the future reservoir bottom in the shallow Sélingué reservoir was carried out only in the proximity of the dam site. In the deeper reservoir Manantali the savanna forest was cleared from an area of about 25 percent, and the tree gallery was cleared along the river. The initial filling of Manantali reservoir was followed by a nearly complete drainage of the reservoir. After refilling some months later, the dissolved oxygen content stayed reasonably high and therefore there was no fish mortality. Also no hydrogen sulphide or a large-scale spread of noxious aquatic plants developed. At present the fish fauna is dominated by cichlids and pelagic species, i.e. at Sélingué by clupeids. and at Manantali by cyprinids (Leptocypris niloticus). In both reservoirs was the genus Citharinus considerably reduced. At Sélingué reservoir disappeared Protopterus annectens. The fish yield estimate is 27 kg/ha/year for Manantali, and 40 kg/ha/year for Sélingué. Both rerservoirs have recorded a slight increase in electrical conductivity. They both serve as a valuable water resource and fish supply for the Sahel region.