Attributes of the seed bank after a century of primary succession on islands in Lake Hjälmaren, Sweden

1 A large number of islands was created when the water table of Lake Hjälmaren, south central Sweden, was lowered between 1882 and 1886. We have complete lists of vascular plant species for 40 of these islands from 1886, 1892, 1903–04, 1927–28 and 1984–85.
2 We have investigated the seed bank on nine of these islands and compared species composition at different soil depths with the species lists from the islands in 1886–1985, and with the present vegetation in the area of seed bank sampling. We have also investigated the distribution in the soil of seeds from species with different ecological attributes, including seed longevity, successional status, seed weight, seed form and species longevity.
3 Seeds in soil samples were allowed to germinate over the course of two summers with an intermediate cold storage. We found 1944 seeds representing 65 taxa. The mean seed density was 84 seeds dm ^–2 .
4 The similarity between the surface soil (0–3 cm) seed bank and the vegetation at the different vegetation analyses increased from 1886 to 1993. The similarity between the present vegetation and the seed bank decreased with increasing soil depth, and the soil at 12–15 cm had no species in common with the present vegetation. Several species now absent from the vegetation were found in the seed bank.
5 Deeply buried seeds came from early successional, annual species with long-term persistent and low-weight seeds, as expected from seed bank theories, but were slightly elongated, which was in contrast to theories. Spherical seeds were associated with the surface soil, as were short-lived and high-weight seeds from late successional, perennial species.     

Primary succession over sixty years on hundred-year old islets in Lake Hjälmaren,Sweden

New islands were created as a result of an artificial lowering of the water table of Lake Hjälmaren in 1882–1886 and the floras of 40 such islands have been monitored ever since plant colonization started in 1886. Data used in this paper are from belt-transects across ten of these islands which were analysed in 1927–29 and reanalysed in 1986. During this 60-year period shading by trees has increased and shading and litter accumulation have affected the ground flora. Succession includes aspects of both convergence and divergence. Species of dry, open sites and shore species have been excluded from the island interiors, making these parts of the islands more similar, but there has simultaneously been an increased variability of vegetation types of shaded, mesic habitats. There have been no consistent trends in beta-diversity between the two surveys. Differences in successional patterns between island interiors and shore habitats are attributed to an outward migration of plant species and to differences in disturbance regimes.     

3,4-Dichloroaniline is detoxified and exported via different pathways in Arabidopsis and soybean

The metabolic fate of [UL-14C]-3,4-dichloroaniline (DCA) was investigated in Arabidopsis root cultures and soybean plants over a 48 h period following treatment via the root media. DCA was rapidly taken up by both species and metabolised, predominantly to N-malonyl-DCA in soybean and N-glucosyl-DCA in Arabidopsis. Synthesis occurred in the roots and the respective conjugates were largely exported into the culture medium, a smaller proportion being retained within the plant tissue. Once conjugated, the DCA metabolites in the medium were not then readily taken up by roots of either species. The difference in the routes of DCA detoxification in the two plants could be explained partly by the relative activities of the respective conjugating enzymes, soybean containing high DCA-N-malonyltransferase activity, while in Arabidopsis DCA-N-glucosyltransferase activity predominated. A pre-treatment of plants with DCA increased DCA-N-malonyltransferase activity in soybean but not in Arabidopsis, indicating differential regulation of this enzyme in the two plant species. This study demonstrates that DCA can undergo two distinct detoxification mechanisms which both lead to the export of conjugated metabolites from roots into the surrounding medium in contrast to the vacuolar deposition more commonly associated with the metabolism of xenobiotics in plants.     

Transit towards floristic homogenization on oceanic islands in the south-eastern Pacific: comparing pre-European and current floras

Aim  To quantify the occurrence of processes of homogenization or differentiation in the vascular flora of six oceanic islands.
Location  Six islands in the south-eastern Pacific drawn from the Desventuradas Archipelago, Easter Island and the Juan Fernández Archipelago.
Methods  Using published floristic studies, we determined the floristic composition of each island at two different stages: (1) pre-European colonization and (2) current flora. We compared changes in the number of shared plants and the floristic similarity among islands for each stage.
Results  The number of plant species doubled from 263 in pre-European flora to 531 species currently. Only three native species became extinct, four natives were translocated among the islands and 271 plant species were introduced from outside. The frequency of plant species shared by two or more islands is higher in the post-European floras than prior to European contact, and the level of floristic similarity between islands increased slightly.
Main conclusions  Considering the low naturalization rate of alien plants, the small number of extinctions and the meagre increase in floristic similarity, these islands are undergoing a slow process of floristic homogenization.     

A global comparison of plant invasions on oceanic islands

Oceanic islands have long been considered to be particularly vulnerable to biotic invasions, and much research has focused on invasive plants on oceanic islands. However, findings from individual islands have rarely been compared between islands within or between biogeographic regions. We present in this study the most comprehensive, standardized dataset to date on the global distribution of invasive plant species in natural areas of oceanic islands. We compiled lists of moderate (5–25% cover) and dominant (>25% cover) invasive plant species for 30 island groups from four oceanic regions (Atlantic, Caribbean, Pacific, and Western Indian Ocean). To assess consistency of plant behaviour across island groups, we also recorded present but not invasive species in each island group.We tested the importance of different factors discussed in the literature in predicting the number of invasive plant species per island group, including island area and isolation, habitat diversity, native species diversity, and human development. Further we investigated whether particular invasive species are consistently and predictably invasive across island archipelagos or whether island-specific factors are more important than species traits in explaining the invasion success of particular species.We found in total 383 non-native spermatophyte plants that were invasive in natural areas on at least one of the 30 studied island groups, with between 3 and 74 invaders per island group. Of these invaders about 50% (181 species) were dominants or co-dominants of a habitat in at least one island group. An extrapolation from species accumulation curves across the 30 island groups indicates that the total current flora of invasive plants on oceanic islands at latitudes between c. 35°N and 35°S may eventually consist of 500–800 spermatophyte species, with 250–350 of these being dominant invaders in at least one island group. The number of invaders per island group was well predicted by a combination of human development (measured by the gross domestic product (GDP) per capita), habitat diversity (number of habitat types), island age, and oceanic region (87% of variation explained). Island area, latitude, isolation from continents, number of present, non-native species with a known invasion history, and native species richness were not retained as significant factors in the multivariate models.Among 259 invaders present in at least five island groups, only 9 species were dominant invaders in at least 50% of island groups where they were present. Most species were invasive only in one to a few island groups although they were typically present in many more island groups. Consequently, similarity between island groups was low for invader floras but considerably higher for introduced (but not necessarily invasive) species – especially in pairs of island groups that are spatially close or similar in latitude. Hence, for invasive plants of natural areas, biotic homogenization among oceanic islands may be driven by the recurrent deliberate human introduction of the same species to different islands, while post-introduction processes during establishment and spread in natural areas tend to reduce similarity in invader composition between oceanic islands. We discuss a number of possible mechanisms, including time lags, propagule pressure, local biotic and abiotic factors, invader community assembly history, and genotypic differences that may explain the inconsistent performance of particular invasive species in different island groups.     

Strong indication of an extinction‐based saturation of the flora on the Pacific Robinson Crusoe Islands

Oceanic islands are vulnerable ecosystems and their flora has been under pressure since the arrival of the first humans. Human activities and both deliberately and inadvertently introduced biota have had and continue to have a severe impact on island endemic plants. The number of alien plants has increased nearly linearly on many islands, perhaps resulting in extinction‐based saturation of island floras. Here, we provide evidence for such a scenario in Alejandro Selkirk, Robinson Crusoe Islands (Archipelago Juan Fernández, Chile). We compared species richness and species composition of historical vegetation samples from 1917 with recent ones from 2011. Changes in species’ relative occurrence frequency were related to their taxonomic affiliation, dispersal mode, distribution status, and humidity and temperature preferences. While total species richness of vascular plants remained relatively similar, species composition changed significantly. Plants endemic to the Robinson Crusoe Islands declined, exotic species increased substantially within the period of ca. 100 years. Further, the relative occurrence frequency of plants with preferences for very warm and humid climate decreased, while the opposite was found for plants preferring drier and colder environments. Potential drivers responsible for this dramatic shift in the vegetation within only one century might have been the large goat population affecting especially small populations of endemic plants and climatic changes. Taking into account a substantial extinction debt, we expect further shifts in the vegetation of this small oceanic island toward alien plants. This would have significant negative consequences on global biodiversity, considering that island floras contribute substantially to global plant species richness due to their high proportion of endemics.     

PUPPING PHENOLOGY AND DEMOGRAPHY OF HARBOR SEALS (PHOCA VITULINA RICHARDSI)ON TUGIDAK ISLAND, ALASKA

The number of seals on shore at Tugidak Island (Gulf of Alaska) declined 72%–85% between 1976 and 1988 and increased during the 1990s. We compared pupping phenology and the ratio of pupping-period counts to molting-period counts between declining (1976–1979) and increasing (1994–1998) years, and examined the sex/age structure of seals ashore during the 1990s. In the 1970s the onset and peak of pupping occurred 6–18 d later than in the 1990s. Rate of pup abandonment was higher in 1978 than in the 1990s. Between 1994 and 1995, the maximum and mean number of seals ashore increased >50%, largely due to an increase in non-pups. From 1995 to 1998, the sex/age structure of seals ashore was similar among years. We observed three to four times as many seals during the molting period than during the pupping period in the 1970s, whereas similar numbers were ashore during these periods in the 1990s, perhaps reflecting changes in demography and/or haul-out behavior. Changes in pupping phenology and demography may reflect environmental changes, such as food availability, and when monitored in conjunction with population counts, may help us better interpret changes in the number of seals ashore.     

Temporal changes in the island flora at different scales in the archipelago of SW Finland

Question: How have species richness and vegetation patterns changed in a group of islands in the northern Baltic Sea over a 58‐yr period of changing land use and increasing eutrophication? Location: A group of 116 islands, the Brunskär sub‐archipelago, in SW Finland. Methods: A complete survey of vascular plant species performed in 1947–1949 by Skult was repeated by our group using the same methodology in 2005–2007 (historical versus contemporary, respectively). DCAs were performed and total number of species, extinction–colonization rates, species frequency changes and mean Ellenberg indicator values for light, moisture and nitrogen and Eklund indicator values for dependence of human cultural influence were obtained for each island and relevé. Results: Species richness has declined on large islands and increased on small islands. The increase in number of species on small islands is driven by a strong increase in frequency of shore species, which in turn is induced by more productive shores. The decrease in species richness on large islands is related to overgrowth of open semi‐natural habitats after cessation of grazing and other agricultural practices. Conclusions: After the late 1940s, open habitats, which were created and maintained by cattle grazing and other traditional agricultural activities, have declined in favour of woody shrub and forest land. Shores have been stabilized and influenced by the eutrophication of the Baltic Sea, and the vegetation has become more homogeneous. This development, resulting in lower species diversity, poses a challenge for the preservation of biodiversity both on a local and on a landscape level.     

Islands,mainland, and terrestrial fragments: How isolation shapes plant diversity

The fragmentation of natural habitats is a major threat for biodiversity. However, the impact and spatial scale of natural isolation mechanisms leading to species loss, compared to anthropogenic fragmentation, are not clear, mainly due to differences between fragments and islands, such as matrix permeability. We studied a 500 km² Mediterranean region in France, including urban habitat fragments, continuous habitat, and continental‐shelf islands. On the basis of 295 floristic relevés, we built species–area relationships to compare isolation in fragments after urbanization, with continuous habitat and continental‐shelf islands. We assumed either no dispersal, infinite dispersal, or estimated intermediate levels of habitat reachability through graph theory. Isolation mechanisms occurred in fragments but with a lower strength than in near‐shore islands, and most importantly affected perennial plants. Annual plants were less affected, probably due to their smaller size and shorter life cycle. Isolation occurred at landscape level in fragments and at patch level in islands. The amount of reachable habitat (accounting for spatial configuration) explained local species richness in both systems, but the amount of habitat (no consideration of spatial configuration) was already a good predictor. These results suggest an important role of habitat amount around fragments in mitigating the isolation effects observed in near‐shore islands, and the importance of carefully considering different functional groups.     

晋西吕梁山严村流域撂荒地植物群落替中的物种多样性变化

用ＤＣＡ排序研究了黄土丘陵区撂荒地上植物群落演替的趋势和方向,用６种多样性指数分析了物种丰富度、均匀度和综合多样性在演替过程中的变化。随着演替的进展,物种丰富性显著增高,均匀性逐渐降低,综合多样性逐渐升高。     

The ecological impact of oceanic island colonization – a palaeoecological perspective from the Azores

Aim In many cases, human colonization drastically modified the ecosystems of remote oceanic islands before scientists arrived to document the changes. Palaeoecological records before and after human colonization provide insights into the original ecosystems and an assessment of subsequent human impact. We used pollen analysis to compare the impact of 15th century colonization of the Azores with that of natural disturbances such as volcanic eruptions and climate changes. Location Azores archipelago, Atlantic Ocean. Methods Sediment records from three highland sites in the Azores (on the islands of Pico and Flores) were dated radiometrically and analysed palynologically. Pollen taxa were classified as native, endemic or introduced based on comparison with flora lists. Data were statistically zoned and temporal trends identified using detrended correspondence analysis. Results Human colonization of the Azores resulted in rapid, widespread, persistent vegetation changes on a scale unprecedented in the last 2700 years, detectable through the decline of dominant trees, the spread of grasses and fire‐tolerant species, the introduction of exotic plants, evidence for grazing and fire, and changes to soils and moisture availability. During the same period, volcanic eruptions appear to have had more localized impacts on the vegetation, lasting 500–1000 years and favouring endemic taxa. The effect of late Holocene climatic changes on the highland vegetation of the Azores seems to have been minor. Palaeoecological data indicate that at least two plant species went extinct on Pico after human colonization and that some plants regarded as introduced were almost certainly part of the original flora of the islands. Despite a consistent signal of human impact, compositional differences between Juniperus brevifolia communities on Pico and Flores remained after colonization. Main conclusions Human colonization had a greater impact on the pristine vegetation of Pico and Flores than climatic changes and volcanic activity during recent millennia. The similarity between post‐colonization changes on the Azores and other oceanic islands suggests a consistent pattern and scale to historical‐era human impact on otherwise pristine ecosystems. These characteristics could be used to further elaborate biogeographical theory and direct conservation efforts towards species that appear most susceptible to human activity.     

Can We Infer Island Introduction and Naturalization Rates from Inventory Data? Evidence from Introduced Plants in Galapagos

Studies of human-mediated rates of introduction of organisms to islands are few, results fall into two models (linear and exponential), and relationships with human population and activities are obscure. Owing to their late settlement and good scientific record, data from Galapagos may be exceptionally informative. The rate of introduction of vascular plant species to Galapagos has been suggested to be exponential, paralleling the rise in human population. However, detailed reconstruction of growth in numbers of introduced plant species, based on historical and recent records, reveals changes in study criteria over the last two centuries, which obscure the true introduction rate. At first, cultivated species were deliberately excluded from most studies. From the 1960s, naturalized cultivated species were included more consistently, but non-naturalized species were still excluded. From the mid-1980s, the latter were deliberately included. Accidental introductions increased linearly from 1807 (the first records) to the present. Escapes from cultivation show increases in rate around 1906 and in the period 1970–1990, the latter coinciding with the first studies directed at areas affected by human activities. Non-naturalized cultivated species rose abruptly from the late 1980s, as they became deliberately studied. There seems to be no direct link with human population size. Data represent rate of discovery rather than true introduction rate, and the changing overall rate reflects changing botanical interests and research effort. Data from other islands also suggest that linear increases in naturalized plants are the norm. Galapagos data do not permit confident statements about the introduction rate of cultivated species, but suggest that this may depend more on human activities than human population size.     

百山祖常绿阔叶林灌草层物种组成和 分布的10年动态

森林群落的动态受多方面因素的影响。林下的草本植物和木本植物幼苗幼树对于森林的更新及动态具有重要作用。为了解百山祖常绿阔叶林群落下层植物物种组成和分布的动态变化, 我们分析了2003-2013年间百山祖5 ha森林动态样地灌草层物种(包括DBH < 1 cm的木本植物和所有草本植物)在物种重要值、α多样性以及物种分布等方面的变化。结果表明: (1) 10年间, 群落灌草层植物个体数减少了28.7%, 其中草质藤本植物个体数下降比例最大(76.9%), 而乔木的幼苗和幼树下降比例最小(14.4%); (2)整个灌草层的物种数明显下降, 同时乔木的幼苗和幼树及直立灌木的均匀度均下降, 而草本植物均匀度有所上升; (3)除短尾柯(Lithocarpus brevicaudatus)、叶萼山矾(Symplocos phyllocalyx)、显脉野木瓜(Stauntonia conspicua)和福建悬钩子(Rubus fujianensis)等植物重要值逆势上扬外, 直立灌木物种重要值10年间的变化值与其在2003年时的重要值呈显著正相关, 而其余各类生活型植物都显示, 初始重要值越高的物种在10年后下降幅度越大; (4)物种分布与生境类型的相关性变化复杂, 除与山谷生境正相关的物种数量增加外, 研究期间与生境存在显著相关性的物种数都有所减少; (5) 10年来仅陡坡处物种多样性指数没有显著下降。总体来说, 百山祖常绿阔叶林灌草层群落仍未达到顶极状态, 2003-2013年间正缓慢向顶极群落演替。群落内部的种间和种内竞争应是当前群落动态的主导因素。     

Remotely Monitoring Change in Vegetation Cover on the Montebello Islands,Western Australia,in Response to Introduced Rodent Eradication

The Montebello archipelago consists of 218 islands; 80 km from the north-west coast of Western Australia. Before 1912 the islands had a diverse terrestrial fauna. By 1952 several species were locally extinct. Between 1996 and 2011 rodents and cats were eradicated, and 5 mammal and 2 bird species were translocated to the islands. Monitoring of the broader terrestrial ecosystem over time has been limited. We used 20 dry-season Landsat images from 1988 to 2013 and estimation of green fraction cover in nadir photographs taken at 27 sites within the Montebello islands and six sites on Thevenard Island to assess change in vegetation density over time. Analysis of data averaged across the 26-year period suggests that 719 ha out of 2169 ha have increased in vegetation cover by up to 32%, 955 ha have remained stable and 0.6 ha have declined in vegetation cover. Over 492 ha (22%) had no vegetation cover at any time during the period analysed. Chronological clustering analysis identified two breakpoints in the average vegetation cover data occurring in 1997 and 2003, near the beginning and end of the rodent eradication activities. On many islands vegetation cover was declining prior to 1996 but increased after rodents were eradicated from the islands. Data for North West and Trimouille islands were analysed independently because of the potential confounding effect of native fauna being introduced to these islands. Mala (Lagorchestes hirsutus) and Shark Bay mice (Pseudomys fieldi) both appear to suppress native plant recruitment but not to the same degree as introduced rodents. Future research should assess whether the increase in vegetation cover on the Montebello islands is due to an increase in native or introduced plants.     

Use of Plant Strategy Ordination,DCA and ANOVA to elucidate relationships among habitats of Salix planifolia and Salix monticola

Abstract. Detrended Correspondence Analysis (DCA) was used on a data set of 112 species from 14 stands dominated by Salix planifolia and/or S. monticola. Environmental variables were determined which might be important to explain differences in stand vegetation composition, and willow stature (height), in willow carrs in the Rocky Mountain National Park. Correlation of environmental variables with DCA stand scores indicate that peat depth and soil redox potential are highly related to the primary DCA axis. Soils in stands of S. planifolia are more reduced in the early summer and have greater peat depths, than soils in stands of S. monticola. The second DCA axis is related to a water chemistry gradient of hydrogen ion, sulphate, and ammonium concentration. Four environmental variables selected by correlation analysis were regressed on stand scores from the first two DCA axes. Regression coefficients of August soil redox potential and soil water pH were significantly non-zero on the primary DCA axis. In addition to overall vegetation relationships elucidated by DCA, the relationships between willow stature and environmental variables were tested by simultaneous ANOVAs. ANOVA results and DCA ordinations indicate that S. planifolia and S. monticola plants achieve the greatest stature in the least reduced soils, and in shallow peat. Competitive, stress tolerant, and ruderal plant strategy theory applied to the vegetation data produced an arrangement of stands highly correlated to the DCA Axis 1 stand arrangement. These correlations indicate that the primary axis of vegetation structure can be extracted either by species composition, or species morphology/physiology relationships. Concurrent use of DCA and plant strategy theory is suggested as a tool for the prediction of community composition.     

基于叶片解剖性状探究29种植物对热带珊瑚岛的适应策略

为探究植物对热带珊瑚岛高温、强光照、干旱、盐碱等极端环境的适应策略，分别以生长于海南文昌苗圃和移栽至热带珊瑚岛的29种植物为研究对象，对其叶片性状进行测定和比较分析。结果表明，与海南文昌的同种植物相比，热带珊瑚岛的草本植物除干物质含量增加外其他性状均无显著改变；而木本植物的比叶面积显著下降，叶片厚度和叶片干物质含量显著增加，海绵组织更厚且栅栏组织排列更紧密，但气孔长度、气孔密度及气孔面积指数无显著差异。岛上木本植株采取慢速投资-收益的资源获取策略，将更多的资源投入到叶片构建中从而增强逆境下的适应性，并且通过非气孔调节的方式提高光合与储水的潜力以抵御胁迫。栽植于热带珊瑚岛的29种植物能够采用更保守的资源利用策略，较好地适应干旱、强光照等胁迫环境，可用于热带珊瑚岛植被构建。     

Plants and birds on the shore of the Sea of Okhotsk: Balance, crisis, adaptation

Mutual relationships between sea colonial birds and plants on the northern shore of the Sea of Okhotsk are considered. The birds form huge flocks on small islands thus transforming the island landscape and providing specific vegetation at nesting sites. The intensity of ornithogenic load determines different relations between the productivity of plant communities and their species richness. As a rule, flora is depleted. The plants thriving at rookeries react to the intensive action of birds by forming specific ecobiomorphs. On the islands of the northern part of the Sea of Okhotsk tussocks of pine purple grass existing on the majority of nesting colonies are of the highest significance.     

Richness and Composition of Vascular Plants and Cryptogams along a High Elevational Gradient on Buddha Mountain, Central Tibet

We explored patterns of plant species richness and composition along an elevational gradient (4,985?C5,685?m a.s.l.) on Buddha Mountain, 100?km northwest of Lhasa, Tibet. We recorded the presence of plants and lichens in 1-m² quadrats separated by 25-m elevational intervals (174 quadrats in 29 elevational bands) along a vertical transect with a SE aspect. We recorded 143 total species, including 107 angiosperms, 2 gymnosperms, 27 lichens, and 7 mosses. We measured stone cover in each quadrat, and soil pH, C, N and C/N ratio from two randomly located samples collected from 10-cm depth within each band. C, N and C/N decreased with elevation, stoniness increased and soil pH did not change with altitude. We employed detrended correspondence analysis (DCA), canonical correspondence analysis (CCA) and generalized linear models (GLMs) to assess the relationships of species richness and species composition to the environment. The first two axes of the CCA biplot explained 87.7% of total variation in the species-environment relationship, and 27.7% of total variance of species data. The first CCA axis is associated with elevation, while the second axis is related to soil pH and stone cover. We also compared patterns in species richness against expectations from species pools interpolated from the literature. Total species richness was relatively constant between 4,985 and 5,400?m a.s.l. and declined continuously above 5,400?m a.s.l. Similar declining patterns were observed for forbs and graminoids. Cushion plants and lichens abundance exhibited a unimodal relationship with altitude while shrubs declined monotonically. Except for lichens, models derived from our observations and the literature were quite similar in shape. The proportion of the species pool represented in each elevational band increased as a function of elevation for non-vascular plants, but decreased markedly for vascular plants. Thus, vascular plants are more likely to be constrained by dispersal at higher elevations, resulting in more local endemism, while the relatively easily-dispersed high-elevation cryptogams have little local differentiation. Our comparative approach demonstrates that complex scale-dependent differences between life forms may underlie the apparent simplicity of elevational gradients. Furthermore, elevational gradients summarized from distributional notes cannot be assumed to be proxies for elevational gradients on individual mountain slopes.     

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.     

Not across the North Pole: plant migration in the Arctic

? The vascular plant flora of 66 arctic islands was studied to determine whether the islands have been occupied by random long-distance dispersal (LDD) or in a highly structured northward migration pattern via intervening islands as stepping-stones. ? A maximum parsimonious migration model minimizing dispersal distances of 1256 vascular plant taxa was calculated in the framework of network analysis. ? Plant dispersal is not stochastic in the Arctic at the global scale. Inferred mean dispersal distances of the plants occurring on arctic islands are c. 580 km (median 460 km). A LDD across the North Pole could not be inferred in the model and species may be recruited mainly from the nearest mainland or islands. At smaller scales, among adjacent islands, dispersal of vascular plants may be incomplete. Arctic islands do not yet appearto be saturated with species. ? The results suggest that changes in biodiversity in Arctic islands can be more easily predicted at the global scale than at the local scale. Because islands are not yet saturated with species, new colonizations may not necessarily be linked to climate change.