The environmental determinants of the insular butterfly faunas of the British Isles

A multiple regression analysis was performed upon selected environmental variables for a series of islands in the British Isles, to establish their effects upon the size of the butterfly fauna, measured as he number of species regularly breeding, S_B .
So that the data be normally distributed, the regression analyses were performed upon log₁₀ transformed data only, with the data for outliers, mainland Britain and Ireland, the two largest islands, excluded.
Most highly correlated with the number of butterfly species breeding upon an island is the number breeding within a 25 km radius of the nearest point of the mainland, r ²=0.5941, followed by the correlations with the latitude of the mid-point of the island, r ²=0.5541, the number of plant species comprising the island Hora, r ²=0.5225, and the distance separating the island from the mainland, r ²=0.4514.
A partial correlation analysis confirms the importance of the parameters distance separating the island from the mainland, D ₁, and the size of the faunal source S _F , and rejects the importance of the size of the flora and the latitude of the island. This is further confirmed by the results of a step-wise regression analysis, the two variables D ₁ and S_F accounting for 66% of the variation of the butterfly fauna.
If an alternative measure of isolation, D ₂, which allows for the geographical clumping of islands, is combined with the variable S_F , then 69% of the variation of the butterfly fauna is accounted for.     

Insect biogeography in the south-western Sea of Korea with comments on the insect fauna of Kwanmae Island

The insect species richness of each island in the south‐western Sea of Korea was considered on the background of the equilibrium theory. The species number of insects on Kwanmae Island in the present study (140 species) was much higher compared with a previous survey. Based on a literature survey of island biota surveys published in the 1980s, of 47 islands, the lowest species number (12 species) was on Kwanmae Island, and the most diverse insect fauna (254 species) was on Baekryong Island. The mean species number of surveyed islands was approximately 54 (53.96 with a standard deviation of 46.95). The median species number was 38 with a skew of +2.56. Insects, including the orders Odonanta, Orthoptera, Hemiptera, Lepidoptera, Coleoptera, Hymenoptera and Diptera, occurred on 32 of the 47 surveyed islands, and had an occurrence rate of more than 0.68. This indicates that these insects are distributed widely on the islands in the south‐western Sea of Korea. The species number showed a significant linear relationship with both area of an island and its distance from the mainland (P < 0.05), with an extremely low determinant coefficient (r² = 0.13 for area vs species number and r² = 0.28 for distance vs species number). Other factors tested in the study failed to show a significant relationship with species number. A multiple‐regression model established using area and distance as independent variables showed significant relationship with species number, with a relatively higher determinant coefficient (r² = 0.70, P < 0.05). We present possible explanations to explain the difference between estimated and observed species number in Kwanmae Island.     

Island biogeography theory explains the genetic diversity of a fragmented rock ptarmigan (Lagopus muta) population

The island biogeography theory is one of the major theories in ecology, and its applicability to natural systems is well documented. The core model of the theory, the equilibrium model of island biogeography, predicts that species diversity on an island is positively related to the size of the island, but negatively related by the island's distance to the mainland. In recent years, ecologists have begun to apply this model when investigating genetic diversity, arguing that genetic and species diversity might be influenced by similar ecological processes. However, most studies have focused on oceanic islands, but knowledge on how the theory applies to islands located on the mainland (e.g., mountain islands, forest islands) is scarce. In this study, we examined how the size and degree of isolation of mountain islands would affect the genetic diversity of an alpine bird, the rock ptarmigan (Lagopus muta). Within our study area, we defined the largest contiguous mountain area as the mainland, while smaller mountains surrounding the mainland were defined as islands. We found that the observed heterozygosity (H_o) was significantly higher, and the inbreeding coefficient (F_is) significantly lower, on the mainland compared to islands. There was a positive significant relationship between the unbiased expected heterozygosity (H_n.b.) and island size (log km²), but a negative significant relationship between H_o and the cost distance to the mainland. Our results are consistent with the equilibrium model of island biogeography and show that the model is well suited for investigating genetic diversity among islands, but also on islands located on the mainland.     

Population and colony structure of the carpenter ant Camponotus floridanus

The colony and population structure of the carpenter ant, Camponotus floridanus, were investigated by multilocus DNA fingerprinting using simple repeat motifs as probes [e.g. (GATA)₄]. The mating frequency of 15 queens was determined by comparing the fingerprint patterns of the queen and 17–33 of her progeny workers. C. floridanus queens are most probably singly mated, i.e. this species is monandrous and monogynous (one queen per colony). C. floridanus occurs in all counties of mainland Florida and also inhabits most of the Key islands in the southern part of Florida. We tested whether the two mainland populations and the island populations are genetically isolated. Wright's F_ST and Nei's D-value of genetic distance were calculated from intercolonial bandsharing-coefficients. The population of C. floridanus is substructured (F_ST= 0.19 ± 0.09) and the highest degree of genetic distance was found between one of the mainland populations and the island populations (D= 0.35). Our fingerprinting technique could successfully be transferred to 12 other Camponotus species and here also revealed sufficient variability to analyse the genetic structure. In three of these species (C. ligniperdus, C. herculeanus and C. gigas) we could determine the mating frequency of the queen in one or two colonies, respectively.     

Nestedness in island faunas: novel insights into island biogeography through butterfly community profiles of colonization ability and migration capacity

Aim To relate variation in the migration capacity and colonization ability of island communities to island geography and species island occupancy. Location Islands off mainland Britain and Ireland. Methods Mean migration (transfer) capacity and colonization (establishment) ability (ecological indices), indexed from 12 ecological variables for 56 butterfly species living on 103 islands, were related to species nestedness, island and mainland source geography and indices using linear regression models, RLQ analysis and fourth‐corner analysis. Random creation of faunas from source species, rank correlation and rank regression were used to examine differences between island and source ecological indices, and relationships to island geography. Results Island butterfly faunas are highly nested. The two ecological indices related closely to island occupancy, nestedness rank of species, island richness and geography. The key variables related to migration capacity were island area and isolation; for colonization ability they were area, isolation and longitude. Compared with colonization ability, migration capacity was found to correlate more strongly with island species occupancy and species richness. For island faunas, the means for both ecological indices decreased, and variation increased, with increasing island species richness. Mean colonization ability and migration capacity values were significantly higher for island faunas than for mainland source faunas, but these differences decreased with island latitude. Main conclusions The nested pattern of butterfly species on islands off mainland Britain and Ireland relates strongly to colonization ability but especially to migration capacity. Differences in colonization ability among species are most obvious for large, topographically varied islands. Generalists with abundant multiple resources and greater migration capacity are found on all islands, whereas specialists are restricted to large islands with varied and long‐lived biotopes, and islands close to shore. The inference is that source–sink dynamics dominate butterfly distributions on British and Irish islands; species are capable of dispersing to new areas, but, with the exception of large and northern islands, facilities (resources) for permanent colonization are limited. The pattern of colonization ability and migration capacity is likely to be repeated for mainland areas, where such indices should provide useful independent measures for assessing the conservation status of faunas within spatial units.     

Crossover of a high-spin (S=7/2, 3/2) to a low-spin (S=1/2) iron-selenium cluster in FA and FB of photosystem I on rebinding of PsaC onto P700-FX cores

 PsaC is a tightly bound ferredoxin in the Photosystem I (PS I) reaction center which contains two [4Fe-4S] clusters named F_A and F_B. We recently proposed that the mixed-ligand F_B cluster in C14D_PsaC and the mixed-ligand F_A cluster in C51D_PsaC exist in a spin state of S=3/2, and that a spin state crossover to S=1/2 occurs when the PsaC mutants are rebound onto P700-F_X cores. Since EPR signals from a highly rhombic S=3/2 spin state can be difficult to study, wild-type PsaC was reconstituted with iron and selenium to introduce an easily detected S=7/2 spin state similar to that shown for Clostridial ferredoxin. When the unbound [4Fe-4Se] PsaC was chemically reduced, a sharp derivative resonance was found at g=5.171 attributed to the excited ±3/2 doublet from an S=7/2 spin multiplet. An additional peak was found at g=5.616 attributed to the superimposed ±1/2 and ±3/2 doublets from a highly rhombic S=3/2 spin multiplet, and an axial set of resonances found around g=2.0 attributed, in part, to a classical S=1/2 spin state. When the [4Fe-4Se] PsaC was rebound onto P700-F_X cores, the spin population derived from the S=7/2 and 3/2 spin states was negligible. Illumination of the rebuilt PS I complex at 15 K resulted in two rhombic sets of resonances, one with g values of 2.043, 1.941 and 1.854, diagnostic of F_A, and the other with g values of 2.067, 1.941 and 1.878, diagnostic of F_B. Chemical reduction with sodium dithionite at pH 10.5 or photoaccumulation by freezing during illumination resulted in a set of resonances with g values of 2.046, 1.938, 1.920 and 1.883, characteristic of a spin-coupled F_A ^–/F_B ^– pair. The spin state crossover in this iron chalcogenide cluster is the first known to be induced by protein-protein association and reinforces the hypothesis that an S=3/2 to 1/2 crossover occurs in the PS I-rebound mutants C14D_PsaC and C51D_PsaC. Received: 6 August 1996 / Accepted: 28 December 1996     

Parasites and the island syndrome: the colonization of the western Mediterranean islands by Heligmosomoides polygyrus (Dujardin, 1845)

Aim Populations of free‐living vertebrates on islands frequently differ from their mainland counterparts by a series of changes in morphometric, life‐history, behavioural, physiological and genetic traits, collectively referred to as the ‘island syndrome’. It is not known, however, whether the ‘island syndrome’ also affects parasitic organisms. The present study establishes the colonization pattern of the Mediterranean islands by the nematode Heligmosomoides polygyrus, a direct and specific parasite of rodent hosts of the Apodemus genus, and evaluates the effects of island colonization by this species on two components of the island syndrome: the loss of genetic diversity and the enlargement of the ecological niche. Location Heligmosomoides polygyrus was sampled on seven western Mediterranean islands ? Corsica, Crete, Elba, Majorca, Minorca, Sardinia and Sicily ? as well as in 20 continental locations covering the Mediterranean basin. Methods The mitochondrial cytochrome b gene (690 base pairs) was sequenced in 166 adult H. polygyrus individuals sampled in the 27 continental and island locations. Phylogenetic reconstructions in distance, parsimony, maximum likelihood and Bayesian posterior probabilities were carried out on the whole cytochrome b gene data set. The levels of nucleotide, haplotype and genetic divergence (Kimura two‐parameter distance estimator) diversities were estimated in each island population and in the various continental lineages. Results Phylogenetic reconstructions show that the mainland origins of H. polygyrus were continental Spain for the Balearic Islands (Majorca, Minorca), northern Italy for the Tyrrhenian Islands (Corsica, Sardinia, Elba), southern Italy for Sicily, and the Balkan region for Crete. A comparison of island H. polygyrus populations with their mainland source populations revealed two characteristic components of the island syndrome in this parasite. First, island H. polygyrus populations display a significant loss of genetic diversity, which is related (r² = 0.73) to the distance separating the island from the mainland source region. Second, H. polygyrus exhibits a niche enlargement following insularization. Indeed, H. polygyrus in Corsica is present in both A. sylvaticus and Mus musculus domesticus, while mainland H. polygyrus populations are present exclusively in Apodemus hosts. Main conclusions Our results show that H. polygyrus has undergone a loss of genetic diversity and a niche (host) enlargement following colonization of the western Mediterranean islands. To our knowledge, this study provides the first evidence for components of the ‘island syndrome’ in a parasitic nematode species.     

Mammalian species richness on islands on the Sunda Shelf,Southeast Asia

Summary A rich mammalian fauna is found on islands that lie on the Sunda Shelf, a continental shelf extending from Vietnam to Borneo and Java that was periodically exposed as dry land during the Pleistocene. The correlation between log of island area and number of species is high (r ²=0.94); the slope of the curve is moderate (z=0.235). Distance from small islands to source areas (=Borneo, Sumatra, and the Malay Peninsula) does not appear to affect species richness, nor does depth of water to source area (a measure of isolation time). A species-area curve for forest reserves of varying sizes on the Malay Peninsula has a low slope (z=0.104); comparison of the mainlaind and island curves indicates that decreasing island area is strongly correlated with increased extinction. Extinction has left reduced but ecologically balanced sets of species on all islands, except that carnivores are under-represented on all but the largest islands. Initial body size and rarity appear to play a significant role in determining the probability of extinction of individual species.     

Butterflies of European islands: the implications of the geography and ecology of rarity and endemicity for conservation

Depending on their faunal content islands can function as important ‘vehicles’ for conservation. In this study, we examine data on 440 butterfly species over 564 European islands in 10 island groups. To determine the status of the butterfly fauna, we have adopted two approaches, island-focused and species-focused, examined using principal components analysis and regression modelling. In the former, we relate species richness, rarity and endemicity to island geography (area, elevation, isolation and location in latitude and longitude); in the latter, species occurrence on islands is examined in relation to distribution, range, range boundaries, and altitudinal limits on the continent as well as species’ ecology (number of host plants) and morphology (wing expanse). Species on islands are also assessed for their status on the continental mainland, their distributional dynamics (extinctions, distribution changes) and conservation status (Red Data Book, European Habitat Directive, Species of European Conservation Concern and Bern Convention listing. Unexpectedly, we find that a large fraction of the European butterfly species is found on the islands (63.4%; 59% on small islands) comprising some 6.2% of the land area of Europe. Although species occurring on the islands tend, on the whole, to have lower conservation status and are not declining over Europe, 45 species are endemics restricted to the islands. Species richness shows only a weak locational pattern and is related as expected to isolation from the continental source and island area; but, both rarity and endemicity have distinctive geographical bias to southern Europe, on islands now under increasing pressure from climate change and increasingly intensive human exploitation. The vulnerability of species on islands is emphasised in the relationship of island occurrence (% occurrence and presence/absence of species on any island) with continental distributions. A large proportion of the variation (84%) is accounted by continental distribution, the southern range limit and lower altitudinal limit. Most species (69%) occur on very few islands (<5%). In view of ongoing species dynamics on islands, migrations and extinctions of species, island repositories of species depend in large part on conservation of butterflies at continental sources. The unique faunas and rare species on islands also depend on appropriate concern being given to the island faunas. Conservation of European islands is thus a two-way process, sustaining sources and conserving island refuges. Residuals from the regressions (islands with more or fewer species, rare and endemic species; species occurring more or less frequently than expected on islands) provide warning signals of regions and islands deserving immediate attention.     

Contemporary geography dominates butterfly diversity gradients within the Aegean archipelago (Lepidoptera: Papilionoidea, Hesperioidea)

Aim We compare the influence of contemporary geography and historical influences on butterfly diversity for islands in the Aegean archipelago. Location The Aegean archipelago (Greece) and two islands (Cyprus and Megisti) in the Levantine Sea. Methods Thirty‐one islands were examined. Data are taken from own surveys (Coutsis and Olivier) and from the literature. Stepwise multiple regression is used to determine relationships between species richness, frequency, rarity and endemicity against potential geographical predictors. Stepwise logit regression is used to determine geographical predictors of species incidence on islands. Inter‐island and inter‐species associations have been examined using multivariate ordination and clustering techniques. Results The Aegean butterfly fauna is characterized by decreasing diversity and rarity, and increasing homogeneity, from the periphery to the present geographical centre of the archipelago (Cyclades). Diversity and rarity are shown to relate closely to species richness, and species richness, in turn, is largely explained by contemporary geography, particularly the degree of isolation from the nearest mainland sources of Greece or Turkey, and island dimensions. Islands towards the centre of the archipelago are characterized by a group of mobile species (n ≥ 20 species) with extensive ranges across Europe; species that would have recolonized Santorini (Thira) following the VI6 eruption there c. 1630 bc . Endemic components, indicative of autochthonous evolutionary events, are few (5% of species are endemic) compared to known sedentary organisms (molluscs and isopods), but exceed those for more mobile animals (i.e. birds); their distribution is mainly confined to large isolated islands along the Aegean arc (i.e. Kriti) and in the Dodecanese group. Main conclusions Contemporary geography, i.e. processes currently operating in ecological time, dominates butterfly diversity gradients (species richness, frequency, rarity and incidence) in the archipelago. Two reasons are suggested to account for the lack of endemism and the pattern of decreasing diversity into the Cyclades. First, relict butterfly elements may have become extinct on all but a few larger islands, particularly from environmental changes since the Neolithic (fire and overgrazing). Second, colonization from the continental landmasses is ongoing with more mobile species transferring even to the most isolated islands.     

The genetic heterogeneity of deer mouse populations (Peromyscus maniculatus) in an insular landscape

A survey of the genetic variability in deer mouse populations was performed using specimens collected from six different islands on a lake covering approximately 50 km². Random amplified polymorphic DNA (RAPD) was used to measure the extent of the genetic differences in this insular system. An analysis of molecular variance (AMOVA) revealed that populations are clearly separated at this microgeographic scale (F _st = 0.13863; P < 0.001). The homogeneity of molecular variance test (HOMOVA) indicated that within-population levels vary greatly (B _p = 0.76831; P < 0.001). The within-population molecular variance was found to be mainly correlated with the accessibility of the islands, computed as the inverse of the geographic distance separating an island from the lakeshore (r = 0.916; P < 0.003). Received: March 5, 1999 / Accepted: July 16, 1999     

Adaptive divergence despite strong genetic drift: genomic analysis of the evolutionary mechanisms causing genetic differentiation in the island fox (Urocyon littoralis)

The evolutionary mechanisms generating the tremendous biodiversity of islands have long fascinated evolutionary biologists. Genetic drift and divergent selection are predicted to be strong on islands and both could drive population divergence and speciation. Alternatively, strong genetic drift may preclude adaptation. We conducted a genomic analysis to test the roles of genetic drift and divergent selection in causing genetic differentiation among populations of the island fox (Urocyon littoralis). This species consists of six subspecies, each of which occupies a different California Channel Island. Analysis of 5293 SNP loci generated using Restriction‐site Associated DNA (RAD) sequencing found support for genetic drift as the dominant evolutionary mechanism driving population divergence among island fox populations. In particular, populations had exceptionally low genetic variation, small N_e (range = 2.1–89.7; median = 19.4), and significant genetic signatures of bottlenecks. Moreover, islands with the lowest genetic variation (and, by inference, the strongest historical genetic drift) were most genetically differentiated from mainland grey foxes, and vice versa, indicating genetic drift drives genome‐wide divergence. Nonetheless, outlier tests identified 3.6–6.6% of loci as high F_ST outliers, suggesting that despite strong genetic drift, divergent selection contributes to population divergence. Patterns of similarity among populations based on high F_ST outliers mirrored patterns based on morphology, providing additional evidence that outliers reflect adaptive divergence. Extremely low genetic variation and small N_e in some island fox populations, particularly on San Nicolas Island, suggest that they may be vulnerable to fixation of deleterious alleles, decreased fitness and reduced adaptive potential.     

Rule reversal: Ecogeographical patterns of body size variation in the common treeshrew (Mammalia,Scandentia)

There are a number of ecogeographical “rules” that describe patterns of geographical variation among organisms. The island rule predicts that populations of larger mammals on islands evolve smaller mean body size than their mainland counterparts, whereas smaller‐bodied mammals evolve larger size. Bergmann's rule predicts that populations of a species in colder climates (generally at higher latitudes) have larger mean body sizes than conspecifics in warmer climates (at lower latitudes). These two rules are rarely tested together and neither has been rigorously tested in treeshrews, a clade of small‐bodied mammals in their own order (Scandentia) broadly distributed in mainland Southeast Asia and on islands throughout much of the Sunda Shelf. The common treeshrew, Tupaia glis, is an excellent candidate for study and was used to test these two rules simultaneously for the first time in treeshrews. This species is distributed on the Malay Peninsula and several offshore islands east, west, and south of the mainland. Using craniodental dimensions as a proxy for body size, we investigated how island size, distance from the mainland, and maximum sea depth between the mainland and the islands relate to body size of 13 insular T. glis populations while also controlling for latitude and correlation among variables. We found a strong negative effect of latitude on body size in the common treeshrew, indicating the inverse of Bergmann's rule. We did not detect any overall difference in body size between the island and mainland populations. However, there was an effect of island area and maximum sea depth on body size among island populations. Although there is a strong latitudinal effect on body size, neither Bergmann's rule nor the island rule applies to the common treeshrew. The results of our analyses demonstrate the necessity of assessing multiple variables simultaneously in studies of ecogeographical rules.     

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.     

Effects of population size and isolation on reproductive output in Aquilegia canadensis (Ranunculaceae)

It is generally expected that small, isolated populations will suffer reduced fitness due to inbreeding, yet few studies have investigated the relation between population characteristics, inbreeding and fitness. Among Ontario populations of the short‐lived, perennial plant Aquilegia canadensis, large populations (N>90 flowering plants) outcross twice as frequently as small populations (N=30–40), and inbreeding depression is extremely strong. We tested the prediction that reproductive output, a major component of population fitness, should be positively associated with population size. Data from a survey of 33 populations located on small islands in the St. Lawrence River, Canada and 23 populations on adjacent mainland areas supported this prediction. Population size correlated positively with reproductive output, measured as the number of seedlings produced per plant in 1995 (average r=+0.39 pooled P=0.019), and the number of fruits per plant in 1997 (r=+0.30, P=0.056). We also tested the prediction that fitness should decline with increasing spatial isolation between populations by measuring the distance separating all island populations. However, reproductive output did not correlate with isolation in either year. We compared island and mainland populations to test the prediction that reproductive output should be lower for populations on small islands than those occurring in more continuous mainland habitat. In contrast to our predictions, island populations exhibited, if anything, higher reproductive output than mainland populations. We also found no support for the prediction that the positive association between population size and reproductive output should be stronger for presumably isolated populations on small islands than for those on adjacent mainland areas. While the mechanisms underlying the association between population size and fitness are impossible to identify with correlations alone, our results are consistent with the hypothesis that inbreeding can significantly reduce the fitness of natural populations.     

Factors influencing vascular plant diversity on 22 islands off the coast of eastern North America

Aim The influence of physiographic and historical factors on species richness of native and non‐native vascular plants on 22 coastal islands was examined. Location Islands off the coast of north‐eastern USA and south‐eastern Canada between 41° and 45° N latitude were studied. Island size ranges from 3 to 26,668 ha. All islands were deglaciated between 15,000 and 11,000 yr bp ; all but the four New Brunswick islands were attached to the mainland until rising sea level isolated them between 14,000 and 3800 yr bp . Methods Island species richness was determined from floras compiled or revised since 1969. Simple and multiple regression and rank correlation analysis were employed to assess the relative influence of independent variables on species richness. Potential predictors included island area, latitude, elevation, distance from the mainland, distance from the nearest larger island, number of soil types, years since isolation, years since deglaciation, and human population density. Results Native vascular plant species richness for the 22 islands in this study is influenced most strongly by island area, latitude, and distance from the nearest larger island; richness increases with island area, but decreases with latitude and distance from the nearest larger island as hypothesized. That a similar model employing distance from the mainland does not meet the critical value of P confirms the importance of the stepping‐stone effect. Habitat diversity as measured by number of soil types is also an important predictor of native plant species richness, but at least half of its influence can be attributed to island area, with which it is correlated. Two historical factors, years since deglaciation and years since isolation, also appear to be highly correlated with native species richness, but their influence cannot be separated from that of latitude for the present sample size. Non‐native vascular plant species richness is influenced primarily by island area and present‐day human population density, although human population density may be a surrogate for the cumulative effect of several centuries of anthropogenic impacts related to agriculture, hunting, fishing, whaling, tourism, and residential development. Very high densities of ground‐nesting pelagic birds may account for the high percentage of non‐native species on several small northern islands. Main conclusions Many of the principles of island biogeography that have been applied to oceanic islands apply equally to the 22 islands in this study. Native vascular plant species richness for these islands is strongly influenced by physiographic factors. Influence of two historical factors, years since deglaciation and years since isolation, cannot be assessed with the present sample size. Non‐native vascular plant species richness is influenced by island area as well as by human population density; human population density may be a surrogate for other anthropogenic impacts.     

Population genetic structure of island and mainland populations of the quokka, Setonix brachyurus (Macropodidae): a comparison of AFLP and microsatellite markers

Translocation and reintroduction are important tools for the conservation or recovery of species threatened with extinction in the wild. However, an understanding of the potential genetic consequences of mixing populations requires an understanding of the genetic variation within, and similarities among, donor and recipient populations. Genetic diversity was measured using two independent marker systems (microsatellites and AFLPs) for one island and four small remnant mainland populations of Setonix brachyurus, a threatened medium sized macropod restricted to fragmented habitat remnants and two off-shore islands in southwest Australia. Microsatellite diversity in the island population (R _s = 3.2, H _e = 71%) was similar to, or greater than, all mainland populations (R _s = 2.1–3.9, H _e = 34-71%). In contrast, AFLP diversity was significantly lower in the island population (PPL = 20.5; H _j = 0.118) compared to all mainland populations (mean PPL = 79.5–89.7; mean H _j = 0.23–0.29). Microsatellites differentiated all (mainland and island) populations from each other. However, AFLP only differentiated the island population from the mainland populations—all mainland populations were not significantly differentiated from each other for this marker. Given a known time since isolation of the island population from the mainland (6,000 years ago), and an overall more conservative rate of evolution of AFLP markers, our results are consistent with mainland populations fragmenting thousands of years ago (but <6,000 years), probably as a consequence of reduced rainfall and the constriction of the preferred mesic habitat of quokkas. Our results also support a recent history of severe population bottlenecks in mainland populations, and a long history of bottlenecks of the island population, but reflect a recent explosion in numbers since European occupation of the island. Our results indicate that translocation of island populations to supplement mainland populations would introduce genetically markedly differentiated, and possibly maladapted, individuals.     

Small mammals on Massachusetts islands: the use of probability functions in clarifying biogeographic relationships

Summary Thirty-three insular small mammal communities along the coast of Massachusetts (USA) were surveyed to investigate the biogeographic relationships of the insular communities and to examine the distribution patterns of individual species. Nine species of terrestrial small mammals were observed in the total insular fauna, whereas thirteen occurred on the mainland. The species-area relation yielded a z value of 0.06, which is the lowest value yet reported for insular mammal communities.Multiple logistic regression was used to calculate probability functions for each species in order to identify variables potentially important in determining a species' occurrence on islands and to estimate probabilities of occurrence on islands. Statistically significant and ecologically interpretable functions were obtained for all but one species. Occurrence on islands was positively related to increasing island size in four species and to decreasing island isolation in four species. The extremely low z value, negative correlations of species number with isolation variables, and the inclusion of an isolation variable in the logistic functions of four species indicated that immigration was an important determinant of small mammal occurrence on these islands. There was a positive relationship between population density and number of islands occupied.Logistic regression has several advantages over linear discriminant function analysis, and we suggest that it may be useful in other ecological studies and in the preservation of endangered species.     

Genetic diversity in micronesian island populations of the tropical tree campnosperma brevipetiolata (anacardiaceae)

Tropical plant species have been the focus of considerable attention in regard to their potential economic and social importance in the face of rapidly diminishing biodiversity in the tropics. Pacific Island species represent an even more fragile resource because different island populations are widely scattered and overall population sizes are small. We examined the distribution of genetic variation in Campnosperma brevipetiolata (Anacardiaceae), an upland rainforest tree species that is of potential use for both lumber and reforestation efforts in the Federated States of Micronesia (FSM). Seeds were collected from multiple populations on four island groups in the Caroline Islands (Kosrae, Pohnpei, and Yap in the FSM; and the Republic of Palau) and subjected to an electrophoretic analysis involving four polymorphic genetic loci. We hypothesized that variation on these islands would decrease with increasing distance from the presumed Indo-Malayan source of these island floras. Indeed, we found a trend of decreasing variation from west to east indicated by the mean number of alleles per locus (A = 1.50-1.33), effective number of alleles per locus (A_e = 1.14-1.12) and mean genetic diversity (H_e = 0.123-0.107). We also found little genetic differentiation among the islands (F_pt = 0.174) and among subpopulations within islands (F_sp = 0.047), indicating that either there are high levels of gene flow among the islands by seed dispersal or that these populations have not been established long enough for divergence to have occurred. The lack of divergence among islands observed for Campnosperma brevipetiolata suggests that germplasm sampled from any one island population would be a suitable starting point for plant breeding or reforestation efforts.     

Intercolony variations in movement patterns and foraging behaviors among herring gulls (Larus argentatus) breeding in the eastern Wadden Sea

Herring gulls (Larus argentatus) are opportunistic predators that prefer to forage in the intertidal zone, but an increasing degree of terrestrial foraging has recently been observed. We therefore aimed to analyze the factors influencing foraging behavior and diet composition in the German Wadden Sea. Gulls from three breeding colonies on islands at different distances from the mainland were equipped with GPS data loggers during the incubation seasons in 2012–2015. Logger data were analyzed for 37 individuals, including 1,115 foraging trips. Herring gulls breeding on the island furthest from the mainland had shorter trips (mean total distance = 12.3 km; mean maximum distance = 4.2 km) and preferred to feed on the tidal flats close to the colony, mainly feeding on common cockles (Cerastoderma edule) and shore crabs (Carcinus maenas). In contrast, herring gulls breeding close to the mainland carried out trips with a mean total distance of 26.7 km (mean maximum distance = 9.2 km). These gulls fed on the neobiotic razor clams (Ensis leei) in the intertidal zone, and a larger proportion of time was spent in distant terrestrial habitats on the mainland, feeding on earthworms. δ¹³C and δ¹⁵N values were higher at the colony furthest from the mainland and confirmed a geographical gradient in foraging strategy. Analyses of logger data, pellets, and stable isotopes revealed that herring gulls preferred to forage in intertidal habitats close to the breeding colony, but shifted to terrestrial habitats on the mainland as the tide rose and during the daytime. Reduced prey availability in the vicinity of the breeding colony might force herring gulls to switch to feed on razor clams in the intertidal zone or to use distant terrestrial habitats. Herring gulls may thus act as an indicator for the state of the intertidal system close to their breeding colony.