A multi-scale test for dispersal filters in an island plant community

Constraints on plant distributions resulting from seed limitation (i.e. dispersal filters) were evaluated on two scales of ecological organization on islands off the coast of British Columbia, Canada. First, island plant communities were separated into groups based on fruit morphology, and patterns in species diversity were compared between fruit‐type groups. Second, abundance patterns in several common fleshy‐fruited, woody angiosperm species were compared to species‐specific patterns in seed dispersal by birds. Results from community‐level analyses showed evidence for dispersal filters. Dry‐fruited species were rare on islands, despite being common on the mainland. Island plant communities were instead dominated by fleshy‐fruited species. Patterns in seed dispersal were consistent with differences in diversity, as birds dispersed thousands of fleshy‐fruited seeds out to islands, while dry fruited species showed no evidence of mainland‐island dispersal. Results from population‐level analyses showed no evidence for dispersal filters. Population sizes of common fleshy‐fruited species were unrelated to island isolation, as were rates of seed dispersal. Therefore, island isolation distances were not large enough to impose constraints on species’ distributions resulting from seed limitation. Rates of seed dispersal were also unrelated to island area. However, several species increased in abundance with island area, indicating post‐dispersal processes also help to shape species distributions. Overall results suggest that seed dispersal processes play an important role in determining the diversity and distribution of plants on islands. At the community‐level, dry‐fruited species were seed limited and island communities were instead dominated by fleshy‐fruited species. At the population‐level, common fleshy‐fruited species were not seed limited and showed few differences in distribution among islands. Therefore, although evidence for dispersal filters was observed, their effects on plant distributions were scale‐dependent.     

Genetic structure of the Canarian palm tree (Phoenix canariensis) at the island scale: does the ‘island within islands’ concept apply to species with high colonisation ability?

• Oceanic islands are dynamic settings that often promote within‐island patterns of strong population differentiation. Species with high colonisation abilities, however, are less likely to be affected by genetic barriers, but island size may impact on species genetic structure regardless of dispersal ability.
• The aim of the present study was to identify the patterns and factors responsible for the structure of genetic diversity at the island scale in Phoenix canariensis, a palm species with high dispersal potential. To this end, we conducted extensive population sampling on the three Canary Islands where the species is more abundant and assessed patterns of genetic variation at eight microsatellite loci, considering different within‐island scales.
• Our analyses revealed significant genetic structure on each of the three islands analysed, but the patterns and level of structure differed greatly among islands. Thus, genetic differentiation fitted an isolation‐by‐distance pattern on islands with high population densities (La Gomera and Gran Canaria), but such a pattern was not found on Tenerife due to strong isolation between colonised areas. In addition, we found a positive correlation between population geographic isolation and fine‐scale genetic structure.
• This study highlights that island size is not necessarily a factor causing strong population differentiation on large islands, whereas high colonisation ability does not always promote genetic connectivity among neighbouring populations. The spatial distribution of populations (i.e. landscape occupancy) can thus be a more important driver of plant genetic structure than other island, or species′ life‐history attributes.

     

Seed Rain Produced by Bats and Birds in Forest Islands in a Neotropical Savanna1

The high degree of isolation of forest “islands” relative to “continental” forested areas creates a naturally fragmented landscape in the savanna ecosystem. Because fragmentation can affect the intensity and quality of biological interactions (e.g., seed dispersal) we examined the abundance and species richness of seed rain produced by birds and bats in three different parts of forest islands (center, edge, and exterior) located at the Estación Biológica del Beni, Bolivia. Despite the fact that we found higher species density of seeds in the seed rain at the center of forest islands, when comparing species richness corrected for observed differences in density, species richness was higher at the edge of islands. The three parts of the islands did not differ in total number of seeds. Three genera (Byrsonima, Ficus, and Piper) contributed the most seeds to the seed rain. We found differences in the abundance of dispersed seeds probably because of the variation related with the disturbance line, where the savanna matrix interacts with the forest islands. Carollia perspicillata, Carollia brevicauda, and Sturnira lilium were the bats that contributed most to seed dispersal within forest islands, and Schistochlamys melanopis and Tyranneutes stolzmanni were the most important birds. The movement of seeds produced by bats and birds within forest islands of the savanna is crucial to assure the continuity of ecological process and dynamics of these forest islands.     

片段化景观中壳斗科植物种子捕食和扩散模式

生境片段化伴随的面积效应和边缘效应, 可改变分散贮食动物的竞争强度、觅食行为以及隐蔽条件, 影响种子捕食和扩散模式。阐明生境片段化对多物种种子捕食和扩散的影响, 对理解片段化生境中的植物更新和生物多样性维持十分重要。该研究在浙江省千岛湖地区的岛屿和大陆上开展了针对6种壳斗科植物的种子捕食和扩散实验, 分析了物种、分散贮食动物相对多度、种子产量、岛屿大小和边缘效应如何共同影响种子命运和种子扩散距离。主要结果: (1)种子命运和扩散距离在物种间存在显著差异; (2)大陆比岛屿有更长的种子留存时间, 小岛种子留存时间最短, 岛屿内部比岛屿边缘有更长的种子留存时间; (3)物种和岛屿大小对种子原地取食率存在交互作用, 白栎(Quercus fabri)种子在大岛上有更高的原地取食率; (4)种子在小岛上有最高的扩散率, 分散贮食动物相对多度对种子扩散后贮藏率有负效应。表明在千岛湖地区, 生境片段化改变了种子捕食和扩散模式, 且面积效应对不同物种的种子捕食和扩散模式产生了不同作用, 从而影响森林群落更新和生物多样性维持。     

Abundance-age-area relationships in an insular plant community

Two processes are thought to generate positive relationships between species richness and island area. The areaper se hypothesis states that larger islands maintain larger populations, which are less susceptible to extinction. The habitat hypothesis states that larger islands contain more habitats, and therefore a greater number of habitat specialists. However, the importance of each mechanism is debated. I tested the areaper se and habitat hypotheses by comparing relationships between plant abundance, age and island area in five shrub species on islands off the coast of British Columbia, Canada. Results showed that two shrub species increased in both abundance and age with island area. The remaining three species showed no differences in abundance and age with island area. Conifer abundances increased with island area, which generated differences in habitat availability. Smaller islands were dominated by open habitat, while larger islands contained both open and forested habitats. Changes in habitat availability with island area could explain patterns in plant abundance and age. The two species that increased in abundance with island area were commonly found in conifer forest on the mainland, and their distributions were consistent with the distribution forest habitat. Positive relationships between plant age and island area in these two species may result from lower survivorship in the open habitat, which dominated small islands. The three species that showed no relationship between abundance and island area are commonly found in open habitat on the mainland, and their island distributions paralleled the availability of open habitat on islands. Similar plant ages on different sized islands may result from their occurrence in open habitat on both large and small islands. Overall results support the habitat hypothesis and indicate that species distributions result from the interaction between habitat affinities and changes in habitat availability with island area.     

Deposition of exotic bird-dispersed seeds into three habitats of a fragmented landscape in the northeastern United States

Dispersal by frugivorous birds facilitates invasion by many exotic plants. We measured the seed rain of ornithochorous native and exotic plants at three habitats of a fragmented landscape of the northeastern United States for 1 year. We studied maple-beech forests, old fields, and abandoned conifer plantations. Across all sites we collected 2,196 ornithochorous seeds, including seeds from six exotic species and 10 native species. The majority (90%) of collected seeds were from exotic species. Seed dispersal was broadly similar among habitats, though seed rain of exotic species was higher in old fields than forested habitats. Seed rain was not strongly influenced by artificial perches for most species. However, seeds of exotic species were more commonly found in traps under an artificial perch in old fields. Seed rains for the exotic Elaeagnus umbellata, Rhamnus cathartica, and Rosa multiflora were positively associated with local density of mature plants. Seed rain of R. cathartica was positively associated with abundance of seedlings but not saplings, suggesting that post-dispersal mortality was important. Seed dispersal of the exotic Lonicera spp. was high in all habitats, accounting for 66% of all seeds collected. With the exception of Lonicera spp., seed rain of common exotic invaders was affected by the abundance of seed sources, and these species might be effectively controlled by elimination of local fruiting plants. Fruits of Lonicera morrowii, which has extensively invaded our area, are apparently a common component in the diet of frugivores.     

Variation in dispersability among mainland and island populations of three wind dispersed plant species

Reduced dispersability of species living on islands relative to mainland has been documented in both plants and animals. One evolutionary scenario explains this trend by strong selection against dispersal, once the species has reached the island, to reduce dispersal out to sea. In this study, we compare the dispersal ability of three wind dispersed plant species (Cirsium arvense, Epilobium angustifolium, and E. hirsutum) from populations on mainland and three islands. Dispersal ability was estimated directly as drop time of diaspores, and indirectly using a morphological measure relating the weight of the diaspore to the size of the pappus (Cirsium) or seed hairs (Epilobium). Positive correlation between the morphological measure of dispersal ability and drop time of diaspores were found for all study species. Dispersal ability varied significantly among mainland and islands, and among species. C. arvense showed a significant reduction in dispersal ability on islands compared to mainland, whereas the reverse was found for the two Epilobium species. Overall Epilobium diaspores had a 2–4 times higher dispersability than C. arvense, indicating that degree of isolation of islands vary among study species. Significant differences in dispersability among plants within populations were detected in all species suggesting that this trait may have a genetic component.     

Introduction of mammalian seed predators and the loss of an endemic flightless bird impair seed dispersal of the New Zealand tree Elaeocarpus dentatus

Understanding the mutualistic services provided by species is critical when considering both the consequences of their loss or the benefits of their reintroduction. Like many other Pacific islands, New Zealand seed dispersal networks have been changed by both significant losses of large frugivorous birds and the introduction of invasive mammals. These changes are particularly concerning when important dispersers remain unidentified. We tested the impact of frugivore declines and invasive seed predators on seed dispersal for an endemic tree, hinau Elaeocarpus dentatus, by comparing seed dispersal and predation rates on the mainland of New Zealand with offshore sanctuary islands with higher bird and lower mammal numbers. We used cameras and seed traps to measure predation and dispersal from the ground and canopy, respectively. We found that canopy fruit handling rates (an index of dispersal quantity) were poor even on island sanctuaries (only 14% of seeds captured below parent trees on islands had passed through a bird), which suggests that hinau may be adapted for ground‐based dispersal by flightless birds. Ground‐based dispersal of hinau was low on the New Zealand mainland compared to sanctuary islands (4% of seeds dispersed on the mainland vs. 76% dispersed on islands), due to low frugivore numbers. A flightless endemic rail (Gallirallus australis) conducted the majority of ground‐based fruit removal on islands. Despite being threatened, this rail is controversial in restoration projects because of its predatory impacts on native fauna. Our study demonstrates the importance of testing which species perform important mutualistic services, rather than simply relying on logical assumptions.     

Aggregated seed dispersal by spider monkeys limits recruitment to clumped patterns in Virola calophylla

The initial spatial pattern of seed deposition influences plant population and community structure, particularly when that pattern persists through recruitment. In a vertebrate‐dispersed rain forest tree, Virola calophylla, we found that spatially aggregated seed deposition strongly influenced the spatial structure of later stages. Seed dispersion was clumped, and seed densities were highest underneath V. calophylla females and the sleeping sites of spider monkeys (Ateles paniscus), the key dispersal agent. Although these site types had the lowest per capita seed‐to‐seedling survival, they had the highest seedling/sapling densities. Conversely, seed and seedling/sapling densities were lowest, and seed survival was highest, at sites of diurnal seed dispersal by spider monkeys. Negative density‐dependent and positive distance‐dependent seed survival thinned seed clumps. Nonetheless, the clumped dispersion at sleeping and parental sites persisted to the seedling/sapling stage because differences in seed deposition were large enough to offset differences in seed survival among these site types.     

Colonization of Sphagnum on land uplift islands in the Baltic Sea: time, area, distance and life history

Aim To test whether species richness of Sphagnum mosses on islands in a land uplift archipelago is related to island age, area or connectivity, and whether the frequency of different species can be predicted by their life history and autecology. Location The northern Stockholm archipelago in the Baltic Sea, east‐central Sweden, with a current land uplift rate of 4.4 mm year^?1. Methods We sampled 17 islands differing in area (0.55–55 ha), height (3.6–18 m, representing c. 800–4000 years of age) and distance from mainland (1.6–41 km). For each Sphagnum patch we measured area, height above sea level, horizontal distance from the shore and shading from vascular plants. Factors affecting island species richness, species frequency and habitats on the islands were tested by stepwise regressions. Species frequency was tested on nine life history and autecological variables, including estimated abundance and spore output on the mainland, habitat preference and distribution. Results We recorded 500 patches of 19 Sphagnum species, distributed in 83 rock pools on 14 islands. Island species richness correlated positively with island area and with degree of shelter by surrounding islands, while distance from the mainland, connectivity, height or age did not add to the model. Species frequency (number of colonized islands and rock pools) was mainly predicted by spore output on the mainland and by habitat preference (swamp forest species were more frequent than others), while spore size, for example, did not add to the model. Species differed in mean height above and horizontal distance from the shore, area of occupied rock pools and in the degree of shading of patches. The mean horizontal distance from the shore and the area of occupied rock pools correlated positively with the normal growth position above the water table among species. Spore capsules were found in only 2% of patches, mostly in the bisexual Sphagnum fimbriatum. Main conclusions The presence of Sphagnum in the Stockholm archipelago seems to be governed by regional spore production and habitat demands. Sphagnum does not appear to be dispersal limited at distances up to 40 km and time spans of centuries. Species with a high regional spore output have had a higher colonization rate, which, together with the rarity of spore capsules on the islands, indicate the mainland as a source for colonization rather than dispersal among islands. Swamp forest species seem more tolerant to the island conditions (summer droughts and some salt spray) than open mire species. The different distances from the sea occupied by the species indicate a slow, continuous succession and species replacement towards the island interior as islands are being uplifted and thus expand in area. This partly explains why larger islands harbour more species. Our results thus support some of the island biogeographical theories related to the species–area relationship.     

Patterns of bird species richness and composition on islands off Arnhem Land,Northern Territory,Australia

The bird faunas of the adjacent Wessel and English Company island chains were sampled at two scales (0.25 ha quadrats and entire islands). Ninety‐six species were recorded from 226 quadrats, with the most frequently recorded species being mistletoebird Dicaeum hirundinaceum, brown honeyeater Lichmera indistincta, silver‐crowned friarbird Philemon argenticeps, bar‐shouldered dove Geopelia humeralis, northern fantail Rhipidura rufiventris and yellow white‐eye Zosterops lutea. At the quadrat scale, vegetation type was a major determinant of the abundance of individual species (and hence species composition), species richness and total bird abundance. Bird species composition and richness at the quadrat scale was also significantly affected by island isolation (particularly the amount of land within 20 km of the island perimeter). Island size had no effect on quadrat‐scale richness or total abundance. However, the abundance of 10 of the 38 most frequently recorded individual species was significantly related to island size, in most cases even when the comparison was restricted to similar habitats. The most striking cases were rufous fantail Rhipidura rufifrons, mangrove golden whistler Pachycephala melanura, brown honeyeater and yellow white‐eye, which were all significantly more abundant on smaller islands. One hundred and seventy‐one species were recorded from the 62 islands sampled. There was a very tight relationship between island size and the number of terrestrial species (73% of deviance explained) and of all species (84% of deviance explained). This relationship was improved (marginally) when isolation was included in the model. Ordination of islands by their terrestrial bird species composition was related to island size and isolation, and suggested an erratic species composition on small islands.     

Edge Effects Act Differentially on Multiple Early Regeneration Stages of a Shade‐tolerant Tree Tapirira mexicana

Forest fragmentation is pervasive in tropical landscapes, and one pathway by which fragmentation may negatively impact populations is via edge effects. Early life‐stages are particularly important for species regeneration as they act as bottlenecks, but how edge effects may act differentially on different life‐stages is unknown. This study evaluated edge effects on multiple early life‐stages of a currently common animal‐dispersed, shade‐tolerant tree Tapirira mexicana (Anacardiaceae). The study was conducted in tropical premontane wet forest fragments in a highly deforested region of Costa Rica. The stages assessed were pre‐dispersal predation, primary dispersal, post‐dispersal predation, secondary dispersal, ex situ germination, in situ seed longevity, first and second year seedling abundance, second year seedling survivorship, and basal diameter growth. Results showed that impacts of edge effects were not equal across stages, but were limited to specific stages and times. One stage which may act as a bottleneck for species regeneration was pre‐dispersal predation. Over 60 percent of the seeds were predated by larvae, and predation was higher near the edge than interior habitat. Seeds lost viability within 10 d in the forest. Germination to first year seedling stage was also lower near edges, but such effect was eliminated within a year after that. Primary dispersal, seedling survivorship, and growth were not affected by proximity to edges, and both secondary dispersal and post‐dispersal predation were rare. This study demonstrates that current population abundance may not guarantee future species persistence and the importance of considering multiple life‐stages for a comprehensive assessment of forest fragmentation effects on species regeneration.     

THE RELEVANCE OF GENE FLOW IN METAPOPULATION DYNAMICS OF AN OCEANIC ISLAND ENDEMIC,OLEA EUROPAEA SUBSP. GUANCHICA

Theoretical and empirical studies suggest that geographical isolation and extinction–recolonization dynamics are two factors causing strong genetic structure in metapopulations, but their consequences in species with high dispersal abilities have not been tested at large scales. Here, we investigated the effect of population age structure and isolation by distance in the patterns of genetic diversity in a wind‐pollinated, zoochorous tree (Olea europaea subsp. guanchica) sporadically affected by volcanic events across the Canarian archipelago. Genetic variation was assessed at six nuclear microsatellites (nDNA) and six chloroplast fragments (cpDNA) in nine subpopulations sampled on four oceanic islands. Subpopulations occurring on more recent substrates were more differentiated than those on older substrates, but within‐subpopulation genetic diversity was not significantly different between age groups for any type of marker. Isolation‐by‐distance differentiation was observed for nDNA but not for cpDNA, in agreement with other metapopulation studies. Contrary to the general trend for island systems, between‐island differentiation was extremely low, and lower than differentiation between subpopulations on the same island. The pollen‐to‐seed ratio was close to one, two orders of magnitude lower than the average estimated for other wind‐pollinated, animal‐dispersed plants. Our results showed that population turnover and geographical isolation increased genetic differentiation relative to an island model at equilibrium, but overall genetic structure was unexpectedly weak for a species distributed among islands. This empirical study shows that extensive gene flow, particularly mediated by seeds, can ameliorate population subdivision resulting from extinction–recolonization dynamics and isolation by distance.     

Edge or dispersal effects – Their relative importance on arthropod densities on small islands

Dispersal behaviour and edge effects are two potential factors determining population densities, and both effects are likely to vary with patch size. However, the relative importance of these two effects may be hard to separate because they may produce similar patterns. Here, we separate these two effects on population densities of seven groups of arthropods on small islands. To separate dispersal behaviour and edge effects, we use the fact that the slope of the density–area relationships (DAR-slope) should change with the absolute rates of dispersal, as would occur in response to island isolation, whereas the edge effect is expected not to be dependent on island isolation. For lycosid spiders, parasitic wasps and possibly herbivorous Homoptera DAR-slopes changed between isolated and non-isolated islands, suggesting dispersal behaviour to be relatively more important for explaining variation in their densities. Other arthropods (ants and Collembola), typically those with a predicted low dispersal among islands, showed similar DAR-slopes between isolated and non-isolated islands consistent with dominant edge effects. For two groups (web spiders and Diptera) the results were inconclusive. We conclude that both migratory processes and edge effects should be considered in the evaluation of patch size and isolation on density–area relationships.     

An invasive tree alters the structure of seed dispersal networks between birds and plants in French Polynesia

Aim We studied how the abundance of the highly invasive fruit‐bearing tree Miconia calvescens DC. influences seed dispersal networks and the foraging patterns of three avian frugivores. Location Tahiti and Moorea, French Polynesia. Methods Our study was conducted at six sites which vary in the abundance of M. calvescens. We used dietary data from three frugivores (two introduced, one endemic) to determine whether patterns of fruit consumption are related to invasive tree abundance. We constructed seed dispersal networks for each island to evaluate how patterns of interaction between frugivores and plants shift at highly invaded sites. Results Two frugivores increased consumption of M. calvescens fruit at highly invaded sites and decreased consumption of other dietary items. The endemic fruit dove, Ptilinopus purpuratus, consumed more native fruit than either of the two introduced frugivores (the red‐vented bulbul, Pycnonotus cafer, and the silvereye, Zosterops lateralis), and introduced frugivores showed a low potential to act as dispersers of native plants. Network patterns on the highly invaded island of Tahiti were dominated by introduced plants and birds, which were responsible for the majority of plant–frugivore interactions. Main conclusions Shifts in the diet of introduced birds, coupled with reduced populations of endemic frugivores, caused differences in properties of the seed dispersal network on the island of Tahiti compared to the less invaded island of Moorea. These results demonstrate that the presence of invasive fruit‐bearing plants and introduced frugivores can alter seed dispersal networks, and that the patterns of alteration depend both on the frugivore community and on the relative abundance of available fruit.     

Distributional patterning of terrestrial herpetofauna on the Wessel and English Company Island groups,northeastern Arnhem Land,Northern Territory,Australia

Forty-four species of terrestrial reptiles and eight species of frogs were recorded from 60 continental islands of the Wessel and English Company groups off northeastern Arnhem Land, Northern Territory. Two gecko species, Oedura rhombifer and Heteronotia binoei, were present on the most islands (34 and 31, respectively), and occurred on islands < 5 ha. In contrast, agamids, pygopodids and varanids were absent from islands < 18 ha, and snakes and frogs were not reported from islands < 240 ha. Island size explained 82% of the variation in species richness for terrestrial reptiles, and 84% of that for lizards. The relationship was less good for (i) groups with generally uncommon species (notably snakes), for which sampling effort explained more variation, and (ii) groups with species which had relatively specific habitat requirements (notably frogs), for which island size and isolation factors were not especially relevant. For most taxonomic groups considered, isolation factors added little to the relationship between species richness and island size. Across all reptiles, larger species were found on fewer islands, and had larger island size thresholds. This relationship broke down with analysis restricted to the single most species-rich family, Scincidae. Only 6 of the 20 most frequently recorded species showed significant variation in abundance among 8 vegetation types sampled by 226 quadrats across 40 islands. The number of species (alpha-diversity) and total abundance of herpetofauna within quadrats was generally unrelated to island size; however, (with analysis restricted to islands on which they occurred) six individual species were significantly more abundant on smaller islands than on larger islands, with no species showing the opposite pattern. The islands’ herpetofauna is largely a relatively depauperate subset of that of the far more complex sandstone massif and escarpment of western Arnhem Land, especially missing species associated with rugged sandstone gorges, riparian areas, open forests, swamps and clay soils. Patterns in species richness and composition are explained by greater range of environments on larger islands allowing better retention of species since isolation and/or richer tallies at the time of isolation. The evidence suggests that there has been relatively little colonization, although at least two gecko species and one varanid may have moved reasonably frequently.     

Contrasting diversity and demographic signals in sympatric narrow‐range endemic shrubs of the south‐west Western Australian semi‐arid zone

Comparative studies of sympatric species that integrate both phylogeographical and population genetic approaches provide insight into how demographic events and life history traits shape adaptive potential and drive species persistence. Such studies are rare for species‐rich and strongly structured environments, especially those of the southern hemisphere. For two sympatric, perennial shrubs of the south‐west Western Australian semi‐arid zone, Grevillea globosa and Mirbelia sp. Bursarioides, we assessed historical and contemporary genetic diversity and structure, demographic processes and ratios of pollen to seed dispersal. Phylogeographical structure was not detected and haplotype networks were star‐like. Number of haplotypes, nucleotide diversity, haplotype diversity, and allelic diversity were statistically significantly lower for G. globosa than for M. sp. Bursarioides. Levels of haplotype divergence and more contemporary genetic divergence and expected heterozygosity were lower for G. globosa than for M. sp. Bursarioides, but differences were not statistically significant. Both species exhibited signals of isolation by distance and low pollen to seed dispersal ratios (5.26:1 and 6.88:1). Grevillea globosa displayed signals of historical and contemporary demographic expansion. Results imply an important role for aspects of seed ecology that impact population demography, as well as direct dispersal and a significant contribution of seed dispersal to genetic connectivity in a semi‐arid landscape.     

Historical biogeography of the endemic Campanulaceae of Crete

Aim The clade Campanulaceae in the Cretan area is rich in endemics, with c. 50% of its species having restricted distributions. These species are analysed in the context of a larger phylogeny of the Campanulaceae. Divergence times are calculated and hypotheses of vicariance and dispersal are tested with the aim of understanding whether Cretan lineages represent remnants of an older continental flora. Location The Cretan area: Crete and the Karpathos Islands (Greece). Methods We obtained chloroplast DNA sequence data from rbcL, atpB and matK genes for 102 ingroup taxa, of which 18 are from the Cretan area, 11 are endemics, and two have disjunct, bi‐regional distributions. We analysed the data using beast , a Bayesian approach that simultaneously infers the phylogeny and divergence times. We calibrated the tree by placing a seed fossil in the phylogeny, and used published age estimates as a prior for the root. Results The phylogenetic reconstruction shows that all Campanula species fall within a well‐supported campanuloid clade; however, Campanula is highly polyphyletic. The Cretan endemics do not form a monophyletic group, and species are scattered throughout the campanuloid clade. Therefore, the Cretan taxa did not evolve following a single vicariance or dispersal event. Most Cretan lineages represent remnants of an older continental flora, with the exception of one clade that radiated in situ after island isolation, and one lineage that appears to have arrived by dispersal. Main conclusions Most Cretan species were present in the islands at the time of their isolation, and very little long‐distance dispersal to Crete and diversification within Crete has occurred since then. Endemism is probably driven by loss of species on the mainland after island isolation. Species on the islands may have been more widespread in the past, but they are now restricted to often inaccessible areas, probably as a result of human pressure.     

Are islands more susceptible to plant invasion than continents? A test using Oxalis pes‐caprae L. in the western Mediterranean

Aim We tested the relative vulnerability of islands to Oxalis pes‐caprae L. invasion compared to mainland regions. Oxalis pes‐caprae is a South African annual geophyte that reproduces via bulbils, and has spread in many Mediterranean and temperate regions of the world where introduced. Our study is one of the first detailed regional analyses of the occurrence and local abundance of a non‐native plant. Methods We conducted an extensive survey (2000 sampling points) to examine local and coarse‐scale patterns in both the occurrence and abundance of O. pes‐caprae on islands and in neighbouring mainland regions of Spain. Location We analysed occurrence (number of samples where present) and abundance (percentage cover) on two Balearic Islands (Menorca and Mallorca) and in two mainland administrative provinces of Spain (Murcia and València). Results Oxalis pes‐caprae occurrence was consistently higher on islands. Occurrence varied among habitats, being the highest in tree groves and the lowest in forests and shrublands. It was never found in these two habitats on the mainland. Mean O. pes‐caprae abundance was greatest in tree groves on the mainland, and in field margins and old fields on the islands. However, in general there were not significant differences in local abundances between island and mainland locations. Main conclusions These findings suggest that local processes (such as the biotic resistance of plant communities) are less important than coarse‐scale phenomena (such as environmental driving forces) in explaining differences in the invasion patterns observed between islands and adjacent mainland regions. We suggest that O. pes‐caprae has occupied a larger proportion of available habitats on islands due to: (1) its strong dependence on domestic animal and human‐mediated dispersal which are probably greater on the islands than in mainland areas, and (2) the smaller area encompassed by islands that, over a comparable period of time, enables a greater proportion of available habitats to be colonized (and hence higher occurrence) than equivalent larger mainland areas.     

Dispersal of fig seeds in the Cook Islands: introduced frugivores are no substitutes for natives

Across the Pacific, island vegetation is altering in response to changes in seed disperser assemblages brought about by extinctions and introductions of birds and other animals. On the Cook Islands in the South Pacific, the Pacific Banyan (Ficus prolixa, Moraceae) is undergoing little if any recruitment, possibly linked to a lack of dispersal agents. On Rarotonga, where F. prolixa is found in semi-urban and agricultural environments, there is no recent recruitment in contrast to the situation on Atiu where the tree is common in native forest. We examined the quality and quantity of seed dispersal offered to F. prolixa by the available frugivores on these islands, comparing the effectiveness of extant native and introduced species. The native Cook Islands fauna, particularly birds and bats, appear to be the most effective seed dispersers of F. prolixa, both in terms of quantity and quality. Whilst these are relatively numerous on Atiu, they rarely visit F. prolixa on Rarotonga. The native Chocolate hermit crab Coenobita cavipes is a previously unreported additional native seed disperser, conferring low quantity, but high quality dispersal. Introduced birds and mammals are the most numerous F. prolixa frugivores on Rarotonga and in non-forest environments on Atiu, but they act mainly as seed predators. Consequently, the losses and rarity of remaining native frugivores have not been compensated for by introduced species on Rarotonga which may be contributing to the absence of recruitment there.