Using genetic techniques to quantify reinvasion,survival and in situ breeding rates during control operations

Determining the origin of individuals caught during a control/eradication programme enables conservation managers to assess the reinvasion rates of their target species and evaluate the level of success of their control methods. We examine how genetic techniques can focus management by distinguishing between hypotheses of ‘reinvasion’ and ‘survivor’, and defining kin groups for invasive stoats (Mustela erminea) on Secretary Island, New Zealand. 205 stoats caught on the island were genotyped at 16 microsatellite loci, along with 40 stoats from the opposing mainland coast, and the age and sex were determined for each individual. Using these data, we compare and combine a variety of genetic techniques including genetic clustering, population assignment and kinship‐based techniques to assess the origin of each stoat. The population history and individual movement could be described in fine detail, with results indicating that both in‐situ survival and breeding, and reinvasion are occurring. Immigration to the island was found to be generally low, apart from in 1 year where around 8 stoats emigrated from the mainland. This increased immigration was probably linked to a stoat population spike on the mainland in that year, caused by a masting event of southern beech forest (Nothofagus sp.) and the subsequent rodent irruption. Our study provides an example of some of the ways genetic analyses can feed directly into informing management practices for invasive species.     

Identifying eradication units in an invasive mammalian pest species

Eradication of invasive species is an important component for species conservation and ecosystem restoration. Success of eradications is dependent on knowledge of population connectivity in order to determine reinvasion pathways, and hence populations requiring simultaneous eradication (eradication units). The common brushtail possum (Trichosurus vulpecula) was introduced into New Zealand from Australia and Tasmania, and now occupies a wide range of habitats across the majority of New Zealand. Possums are one of the most destructive invaders within New Zealand, with extensive control operations occurring throughout the country. Understanding the population connectivity of possums on mainland New Zealand (North and South Islands) will enhance the success of planned eradications. We examined the genetic population structure of invasive possums to identify gene flow, thus reinvasion pathways, between seven populations around Dunedin and on the Otago Peninsula where eradication of possums is occurring. Genetic variation at 12 microsatellite loci was comparable between all sampled populations and exhibited a significant isolation by distance pattern. Bayesian clustering methods supported the existence of two population clusters, indicating the presence of a reinvasion pathway onto the Otago Peninsula from urban areas at the Southern end of the Peninsula. To avoid recolonisation, all possums on the Otago Peninsula should be eradicated simultaneously, with the implementation and ongoing maintenance and monitoring of an urban buffer zone. We recommend pre-eradication genetic analyses be adopted by all pest managers to define appropriate eradication units, thereby maximising eradication success and avoiding costly failures.     

Establishing the eradication unit of Molara Island: a case of study from Sardinia, Italy

Molara is a small island belonging to the Marine protected Area Tavolara—Punta Coda Cavallo, in Sardinia. During 2006–2007, a bio-monitoring program reported a strong presence of the black rat, Rattus rattus, on Molara island. Rat predation has detrimentally affected the unique biodiversity of this island, thus, in 2008 an eradication campaign was conducted. Our eradication protocol included a pre-eradication genetic investigation, using 8 microsatellite loci, on a rat population of Molara as well as on neighbour islands within the Marine Protected Area (MPA). The main goal of this genetic investigation was to establish the correct borders of the eradication unit of Molara island. As several recent eradication campaigns have been unsuccessful, due to incomplete and unstable eradication, we also aimed to assess possible hidden sources of reinvasion. Specimens were also collected during post- eradication monitoring on Molara for genetic screening to establish their origin, and thus validate the effectiveness of our eradication campaign. According to our genetic analysis, within the MPA there are four different eradication units, corresponding to the islands of Molara, Tavolara, Piana and to the Sardinia mainland. Gene flow among these four units is more or less absent. The assignment and clustering tests performed on pre and post-eradication samples seem to indicate that the population of Sardinia mainland is a possible source of re-invasion for the Piana and Molara populations.     

Prioritising islands in the United Kingdom and crown dependencies for the eradication of invasive alien vertebrates and rodent biosecurity

Invasive alien vertebrates (IAVs) pose a significant threat to island biodiversity worldwide, and their removal is an important nature conservation management goal. As methods advance, eradications from larger islands and of multiple species simultaneously are increasingly undertaken. Effective targeting to maximise conservation gain is important given limited resources. We build on existing prioritisation methods and use the islands of the UK and Crown Dependencies (UK) as an example of how vertebrate eradications might be prioritised and invasive-free status maintained through biosecurity. For each of the 9688 UK islands, we assessed ecological importance for native vertebrates and the anticipated impacts of the IAVs present to estimate the benefit of restoration based on the feasibility and sustainability of IAV eradications in relation to island size, human population and risk of unassisted reinvasion by swimming. As reinvasion poses a threat to the long-term benefits of eradication, we incorporated species-specific swimming distances and explored the effects of varying reinvasion probability from risk-averse to higher-risk strategies. The 25 islands that would benefit most from eradications were in Scotland, Northern Ireland and the Channel Islands. Our prioritisation method should be seen as an initial guide to identify islands that might benefit from intervention when faced with a large list of potential sites. Feasibility studies taking account of factors such as interspecific interactions, anthropogenic reinvasion, views of residents or ‘social feasibility’ and cost need to be undertaken before planning any eradication. We prioritised biosecurity for rat-free islands to highlight where comprehensive measures might be most beneficial.     

Genetic and phylogenetic consequences of island biogeography

Abstract.— Island biogeography theory predicts that the number of species on an island should increase with island size and decrease with island distance to the mainland. These predictions are generally well supported in comparative and experimental studies. These ecological, equilibrium predictions arise as a result of colonization and extinction processes. Because colonization and extinction are also important processes in evolution, we develop methods to test evolutionary predictions of island biogeography. We derive a population genetic model of island biogeography that incorporates island colonization, migration of individuals from the mainland, and extinction of island populations. The model provides a means of estimating the rates of migration and extinction from population genetic data. This model predicts that within an island population the distribution of genetic divergences with respect to the mainland source population should be bimodal, with much of the divergence dating to the colonization event. Across islands, this model predicts that populations on large islands should be on average more genetically divergent from mainland source populations than those on small islands. Likewise, populations on distant islands should be more divergent than those on close islands. Published observations of a larger proportion of endemic species on large and distant islands support these predictions.     

Survivors or reinvaders? Using genetic assignment to identify invasive pests following eradication

When new individuals from a pest species are detected following eradication, identifying whether the new individuals are survivors from the eradication attempt, or reinvaders from another population, is important for management practices. Pearl Island (512 ha) in New Zealand was the first island in the world on which simultaneous eradication of all three invasive rat species was attempted. Rats were detected again 9 months after the eradication operation. We use genetic assignment methods to discriminate between survivor and reinvader hypotheses. All rats found on Pearl Island after eradication were likely to be reinvaders from an adjacent population on much larger Stewart Island (174,600 ha), suggesting that rats were swimming to the island at a rate much greater than anticipated, but that the original eradication was successful. Adequate genetic signal was obtained from opportunistically collected samples, making the method feasible for conservation managers with limited time and resources.     

Genetic diversity, geographical range and origin of Bemisia tabaci (Hemiptera: Aleyrodidae) Indian Ocean Ms

The whitefly Bemisia tabaci is a pest vector of begomoviruses on crops worldwide. Bemisia tabaci is composed of a complex of cryptic species which barely interbreed. An exception is the Ms from the South West Indian Ocean (SWIO), which crosses in low proportions with the exotic B. The Ms, together with B and Q is part of the same phylogenetic clad. To infer the genetic structure, the geographical range and putative origin of this putative species, microsatellite data and mitochondrial DNA (cytochrome oxydase I) sequences were analysed on an extensive sample set, including all the islands of the region and samples from mainland Africa. Only B and Ms populations were detected across these islands. The exotic B was found only on the islands of Réunion and Mauritius, whereas the Ms is found on all the SWIO islands. Very high isolation by distance was found for the Ms populations between islands of the SWIO, suggesting a long period of presence in this region. Ms populations from mainland Africa had a higher COI diversity than the Ms of the SWIO islands. This diversity is correlated with size and geological ages of the SWIO islands. The population genetic data obtained are in accordance with an origin of Ms in Africa, followed by its expansion and evolution across the SWIO islands prior to human arrival, confirming the status of Ms as indigenous in the SWIO islands.     

Network theory as a method for developing biosecurity preparedness: Netweork topology of Paratrechina longicornis in the Pacific and within New Zealand

One of the greatest challenges in eradicating pest species is determining when no further individuals remain: terminating the control programme too early means failure to eradicate, whereas continuing for too long can add considerable expense. Since monitoring tools are usually only qualitative and invariably imperfect, there may be considerable uncertainty about when and if eradication has been achieved. However, it is possible to quantify the efficacy of monitoring tools and to use this together with knowledge of the basic ecology of the target pest to robustly quantify the probability of successful eradication over time. Here, I describe one such approach and demonstrate its use in the large-scale eradication of painted apple moth (Teia anartoides) from Auckland, New Zealand. A population model for the production of male moths was used in conjunction with spatially-explicit pheromone trap locations and attraction radii to determine the daily probability of detecting a hypothetical wild population at a particular location. Over time, these probabilities compounded to decrease the likelihood of painted apple moth presence given an ongoing lack of detection. In this way, spatio-temporal risk maps were produced to inform managers and to suggest when eradication had been achieved to a predetermined level of certainty. The model suggested that eradication was likely to have been successful in the main infestation areas by mid 2005, with subsequent catches likely to represent further small incursions, as corroborated by evidence from mitochondrial DNA and stable isotope markers. While it was plausible that a wild population was present in the Otahuhu area in 2005, it was very unlikely that it remained by the end of 2006. Population probability models have potential for much wider use in border biosecurity and establishment of area freedom, particularly in combination with future automated trapping systems.     

Using genetics and Bayesian modelling to evaluate the eradication of stoats (

The New Zealand Department of Conservation recently (May 2008) began a programme to eradicate stoats (Mustela erminea) from Resolution Island (Fiordland, New Zealand) using kill traps. In conjunction with this eradication effort we have the following 3 objectives: (1) to measure the population abundance of stoats prior to trapping using hair tubes and forensic DNA methods; (2) optimise techniques for detecting individual stoats, in order to quantify the probability of stoat persistence given no detections after several months of trapping; and (3) use genetic analyses to identify the possible origins (mainland incursions or in situ breeding) of new stoats captured in a control zone. We present Bayesian modelling techniques used to determine the probability of stoat persistence on the island after the initial population reduction, when individual stoats are no longer captured in traps. We also provide details on an effective level of monitoring and trapping effort required to maintain a comfortable level of confidence that stoats no longer persist on the island. Improving these techniques adds to variety of valuable tools for management of invasive mammal species in a range of natural environments worldwide.     

Population probability models for insect eradication

One of the greatest challenges in eradicating pest species is determining when no further individuals remain: terminating the control programme too early means failure to eradicate, whereas continuing for too long can add considerable expense. Since monitoring tools are usually only qualitative and invariably imperfect, there may be considerable uncertainty about when and if eradication has been achieved. However, it is possible to quantify the efficacy of monitoring tools and to use this together with knowledge of the basic ecology of the target pest to robustly quantify the probability of successful eradication over time. Here, I describe one such approach and demonstrate its use in the large-scale eradication of painted apple moth (Teia anartoides) from Auckland, New Zealand. A population model for the production of male moths was used in conjunction with spatially-explicit pheromone trap locations and attraction radii to determine the daily probability of detecting a hypothetical wild population at a particular location. Over time, these probabilities compounded to decrease the likelihood of painted apple moth presence given an ongoing lack of detection. In this way, spatio-temporal risk maps were produced to inform managers and to suggest when eradication had been achieved to a predetermined level of certainty. The model suggested that eradication was likely to have been successful in the main infestation areas by mid 2005, with subsequent catches likely to represent further small incursions, as corroborated by evidence from mitochondrial DNA and stable isotope markers. While it was plausible that a wild population was present in the Otahuhu area in 2005, it was very unlikely that it remained by the end of 2006. Population probability models have potential for much wider use in border biosecurity and establishment of area freedom, particularly in combination with future automated trapping systems.     

Distribution of S-alleles in island populations of flowering cherry, Prunus lannesiana var. speciosa

We surveyed the distribution of S-alleles in natural island populations of Prunus lannesiana var. speciosa sampled from seven sites on the Izu Peninsula and six Izu islands, Japan. The S-genotypes of sampled individuals were determined by Southern analysis of RFLPs generated by restriction enzyme digestion of genomic DNA, using cDNA of the S-RNase gene as a probe. All individuals were heterozygous, as expected under gametophytic self-incompatibility (GSI). Sixty-three S-alleles were observed in the species, but 12 private to the Izu Peninsula population seemed to be derived from related species, giving a total of 75. The estimated number of S-alleles in each population ranged from 26 to 62, and was inversely correlated with the respective population's distance from the Izu Peninsula, the closest point in the mainland to the islands. This geographical cline in the estimated numbers of S-alleles suggests that gene flow to and from the distant island populations was less frequent, and that the studied species has migrated from the mainland to the Izu islands. The genetic relationship at the S-locus among populations also gave an "isolation by distance" pattern. The genetic differentiation at the S-locus among the populations was very low (F(ST) = 0.014, p < 0.001). The number of S-alleles in the species did not seem to depend on genetic differences associated with population subdivisions. This might be due to the greater effective migration rates of S-alleles, as expected under balancing selection in GSI.     

Utility of island populations in re‐introduction programmes – relationships between Arabian gazelles (Gazella arabica) from the Farasan Archipelago and endangered mainland populations

Understanding local adaptation and population differentiation is vital to the success of re‐introduction initiatives. As other mammals living on islands, Arabian gazelles (G. arabica) show reduced body size on the Farasan archipelago, which we corroborated in this study through morphometric analyses of skulls. In the light of the steep population decline on the Arabian Peninsula – but stable population development on the archipelago – we tested the potential suitability of Farasan gazelles as a source for re‐introductions on the mainland. We therefore investigated genetic differentiation between Farasan and mainland populations using eleven nuclear microsatellite loci and detected a distinct genetic cluster exclusively present on the archipelago, which we inferred to be separated from the mainland cluster for less than 2000 years. About 30% of sampled individuals from Farasan Islands showed assignment to a mainland cluster with signs of ongoing introgression. Analyses using the isolation‐with‐migration model confirmed recent (probably human‐induced) bidirectional exchange of gazelles between mainland and island populations. Hence, the surprisingly uniform island dwarfism most likely reflects phenotypic plasticity, that is, altered morphology as a direct consequence of harsh environmental conditions and resource limitation on the archipelago. Should a further decline of Arabian gazelles on the mainland necessitate restocking in the future, Farasan gazelles may thus become an additional source for captive breeding programmes.     

Population genetic structure of Sorex unguiculatus and Sorex caecutiens (Soricidae, Mammalia) in Hokkaido, based on microsatellite DNA polymorphism

We investigated the genetic structure of Sorex unguiculatus and Sorex caecutiens populations in Hokkaido, Japan, using hypervariable microsatellite DNA markers. We used five microsatellite loci to type 475 S. unguiculatus individuals from 20 localities on the Hokkaido mainland and four localities from each of four offshore islands (and 11 shrews from one locality in southern Sakhalin for a particular analysis). We used six microsatellite loci to type 240 S. caecutiens individuals from 13 localities on the Hokkaido mainland. Genetic variation was high in mainland populations of both species and low in the island populations of S. unguiculatus. Allelic richness and island size were positively correlated for S. unguiculatus, suggesting that genetic drift occurred on those islands due to small population size. In addition, four insular populations of S. unguiculatus were genetically differentiated from the mainland populations, although clear phylogeographic clustering was not confirmed among populations on the Hokkaido mainland for either S. unguiculatus or S. caecutiens. Heterozygosity excess was observed in more than half of the populations including the mainland populations of the two species, suggesting recent bottleneck events in these populations. Population dynamics of the shrews might be explained by a metapopulation scheme. According to autocorrelation analysis, the extent of non-random spatial genetic structure was approximately 100 km. Isolation by distance was observed in S. unguiculatus, but not in S. caecutiens although there is a positive trend. The lack of correlation for S. caecutiens might have been due to small sample size. Thus, no obvious differences in population genetic structure were found between the two species on the Hokkaido mainland in the present study, while previous investigations using mitochondrial DNA sequences inferred that these two species might have rather different biogeographic histories.     

Genetic diversity and population size: island populations of the common shrew, Sorex araneus

Populations of many species are currently being fragmented and reduced by human interactions. These processes will tend to reduce genetic diversity within populations and reduce individual heterozygosities because of genetic drift, inbreeding and reduced migration. Conservation biologists need to know the effect of population size on genetic diversity, as this is likely to influence a population's ability to persist. Island populations represent an ideal natural experiment with which to study this problem. In a study of common shrews (Sorex araneus) on offshore Scottish islands, 497 individuals from 13 islands of different sizes and 6 regions on the mainland were trapped and genotyped at eight microsatellite loci. Previous genetic work had revealed that most of the islands in this study were highly genetically divergent from one another and the mainland. We found that most of the islands exhibited lower genetic diversity than the mainland populations. In the island populations, mean expected heterozygosity, mean observed heterozygosity and mean allelic richness were significantly positively correlated with log island size and log population size, which were estimated using habitat population density data and application of a Geographic Information System.     

Genetic and morphometric differentiation among island populations of two Norops lizards (Reptilia: Sauria: Polychrotidae) on independently colonized islands of the Islas de Bahia (Honduras)

Aim Anole lizards (Reptilia: Sauria: Polychrotidae) display remarkable morphological and genetic differentiation between island populations. Morphological differences between islands are probably due to both adaptive (e.g. differential resource exploitation and intra‐ or interspecific competition) and non‐adaptive differentiation in allopatry. Anoles are well known for their extreme diversity and rapid adaptive speciation on islands. The main aim of this study was to use tests of morphological and genetic differentiation to investigate the population structure and colonization history of islands of the Islas de Bahia, off the coast of Honduras. Location Five populations of Norops bicaorum and Norops lemurinus were sampled, four from islands of the Islas de Bahia and one from the mainland of Honduras. Methods Body size and weight differentiation were measured in order to test for significant differences between sexes and populations. In addition, individuals were genotyped using the amplified fragment length polymorphism technique. Bayesian model‐based and assignment/exclusion methods were used to study genetic differentiation between island and mainland populations and to test colonization hypotheses. Results Assignment tests suggested migration from the mainland to the Cayos Cochinos, and from there independently to both Utila and Roatán, whereas migration between Utila and Roatán was lacking. Migration from the mainland to Utila was inferred, but was much less frequent. Morphologically, individuals from Utila appeared to be significantly different in comparison with all other localities. Significant differentiation between males of Roatán and the mainland was found in body size, whereas no significant difference was detected between the mainland and the Cayos Cochinos. Main conclusions Significant genetic and morphological differentiation was found among populations. A stepping‐stone model for colonization, in combination with an independent migration to Utila and Roatán, was suggested by assignment tests and was compatible with the observed morphological differentiation.     

Impact of geographical isolation on genetic differentiation in insular and mainland populations of Weigela coraeensis (Caprifoliaceae) on Honshu and the Izu Islands

Aim To provide insights into genetic differentiation between insular endemic Weigela coraeensis var. fragrans and its progenitor variety W. coraeensis var. coraeensis, the population genetic structure of both varieties was examined, and factors promoting genetic differentiation between the two taxa were explored. Location The natural range of W. coraeensis (sensu lato) throughout mainland Japan (Honshu) and the Izu Islands. Methods The analysis included 349 and 504 individuals across the mainland (Honshu) and the Izu Islands, respectively, using 10 allozyme and 10 microsatellite loci. The population genetic structure of W. coraeensis was assessed by analysing genetic diversity indices for each population, genetic differentiation among populations, model‐based Bayesian clustering or distance‐based clustering, and bottleneck tests. Results The level of genetic diversity in each of the populations on the Izu Islands was negatively correlated with geographical distance between each island and the mainland. The populations on the mainland and on the Izu Islands were genetically differentiated to a certain extent; however, the microsatellite analyses suggested that gene flow also occurred between the mainland and the islands, and among individual islands. These microsatellite analyses also suggested recent bottlenecks in several populations in both areas. Main conclusions The decrease in genetic diversity throughout the Izu Islands, which correlated with distance to the mainland, Honshu, may be the result of a repeated founder effect occurring at a series of inter‐island colonizations from north to south. The stepping stone‐like configuration of the islands may have played a role in the dispersal of the species. Geographical isolation by sea would effectively result in genetic differentiation of W. coraeensis between mainland Honshu and the Izu Islands, although some gene flow may still occur between Honshu and the northern Izu Islands. The differentiation process of the endemic plants on the Izu Islands is anagenetic but not completed, and the study of these plants will provide insightful knowledge concerning the evolution of insular endemics.     

Genetic diversity and colony structure of Tapinoma melanocephalum on the islands and mainland of South China

Aim

Tapinoma melanocephalum is listed as one of the most important invasive pest species in China. Information regarding the patterns of invasion and effects of geographic isolation on the population genetics of this species is largely lacking.

Location

South China.

Methods

To address this problem, we genotyped 39 colonies (two colonies were collapsed due to genetic similarity) using microsatellite markers and mitochondrial DNA sequencing to compare colony genetic structure of T. melanocephalum on the mainland and islands of South China.

Results

An analysis of the colony genotypes showed that the genetic diversity of the mainland population was slightly higher than that of the island populations but not significantly so. However, the observed heterozygosity on Shangchuan Island (SCD) was significantly lower than that of the other colonies. We also found six haplotypes in 111 mitochondrial DNA COI sequences. The relatedness (r) value between colonies of SCD was 0.410, higher than that of the other populations. The genetic clusters among colonies were not related to geographic locations and exhibited admixture likely due to frequent human‐mediated dispersal associated with trade between the mainland population and the islands. Pairwise F_STs between populations showed differentiation among mainland populations, while SCD displayed high levels of divergence (F_ST > 0.15) from most mainland populations. There was no significant isolation by distance among colonies. Most populations showed signs of a bottleneck effect.

Main conclusions

Our study suggests that there was no significant difference in the genetic diversity among the islands and the mainland; however, the lower genetic diversity, the higher degree of genetic divergence from other colonies, and the higher relatedness among nestmates made the SCD population stand out from all the others.     

Genetic variation and structure of house sparrow populations: is there an island effect?

Population genetic structure and intrapopulation levels of genetic variation have important implications for population dynamics and evolutionary processes. Habitat fragmentation is one of the major threats to biodiversity. It leads to smaller population sizes and reduced gene flow between populations and will thus also affect genetic structure. We use a natural system of island and mainland populations of house sparrows along the coast of Norway to characterize the different population genetic properties of fragmented populations. We genotyped 636 individuals distributed across 14 populations at 15 microsatellite loci. The level of genetic differentiation was estimated using F‐statistics and specially designed Mantel tests were conducted to study the influence of population type (i.e. mainland or island) and geographic distance on the genetic population structure. Furthermore, the effects of population type, population size and latitude on the level of genetic variation within populations were examined. Our results suggest that genetic processes on islands and mainland differed in two important ways. First, the intrapopulation level of genetic variation tended to be lower and the occurrence of population bottlenecks more frequent on islands than the mainland. Second, although the general level of genetic differentiation was low to moderate, it was higher between island populations than between mainland populations. However, differentiation increased in mainland populations somewhat faster with geographical distance. These results suggest that population bottleneck events and genetic drift have been more important in shaping the genetic composition of island populations compared with populations on the mainland. Such knowledge is relevant for a better understanding of evolutionary processes and conservation of threatened populations.     

Phylogeography of the scarab beetle Antitrogus parvulus Britton (Coleoptera: Scarabaeidae) in south-eastern Queensland, Australia

Abstract  This study surveys the population genetic structure of Childers canegrub, Antitrogus parvulus , to elucidate its population dynamics and gene flow. Antitrogus parvulus is a pest of sugarcane in the Bundaberg region and this knowledge can be used to optimise integrated pest management practices. Here, base-pair differences in the cytochrome oxidase II gene ( COII ) were used to characterise haplotypic diversity, infer levels of gene flow, and phylogenetic relationships of alleles and their phylogeographical structure. There were 28 unique haplotypes among the 70 sequenced individuals from the seven locations. All three variance components (among regions, among populations, within populations) are highly significant, with highest genetic diversity among regions and lowest among populations within regions. A positive correlation between migration rates and geographical distance and significant phylogeographical structure between four main geographical regions. The main implication of these findings for pest management is that if a grower can eliminate an existing infestation within a field, then reinvasion will be slow and further outbreaks within that field are unlikely to occur. The low dispersal ability of females also means that any resistance to insecticides that develops is likely to remain localised, but will rapidly become dominant within the affected population.     

Species richness, extinction and immigration rates of vascular plants on islands in the Stockholm archipelago, Sweden, during a century of ceasing management

Survey of eighteen islands in the Stockholm archipelago from 1884–1908 was re-examined during 1996–1998 to investigate how island size, degree of isolation and human impact affect the species number, extinction and immigration rates of terrestrial vascular plants. Cattle grazing and haymaking was widely and commonly in practice on the islands, but became less intensive since the 1920s and ceased during the 1960s. In the first survey species number was positively correlated with area and negatively correlated with distance to mainland. This pattern was similar in the later survey. However, the number of surrounding islands was positively correlated with island specific species number. During the period from the first to the second survey twenty species disappeared and ninety-three appeared as new. The increased abundance of trees, bushes and shade tolerant herbs suggest that this increase in species number is the effect of an ongoing succession in the landscape. The absolute extinction rate was negatively correlated with island size and distance to mainland, while the immigration rate was positively correlated to island size. The extinction rate was negatively correlated with the distance to mainland probably due to an earlier more intense cultural exploitation closer to the mainland. Our measure of human impact, the presence of houses on islands, was not related to terrestrial plant species number or extinction rate. However, there was a trend for the immigration rate to be higher on islands with houses in 1942 than those without, indicating the importance of human activities for dispersal and/or recruitment.