Additive partitioning as a tool for investigating the flora diversity in oceanic archipelagos

This paper introduces the integration of additive partitioning with species—area relationships to island biogeography in order to address the question “How are the pteridophyte and spermatophyte native and endemic flora of different oceanic archipelagos partitioned across islands?”.Species richness data of all endemic species and all native species of pteridophytes and spermatophytes were obtained for the Azores, Canaries and Cape Verde in the Atlantic Ocean and Galápagos, Hawaii and Marquesas in the Pacific Ocean. Additive partitioning of species diversity was used to quantify how much of the total diversity of an oceanic archipelago flora (γ-diversity) is due to (i) the mean species richness of the flora of each island (α-diversity), (ii) the variability in species richness of the floras across islands (β_Nestedness) and (iii) the complementarity in species composition of the floras of different islands (β_Replacement). The analysis was separately performed for the native and endemic pteridophyte and spermatophyte floras.The diversity partitioning of the six archipelagos showed large differences in how the flora of each archipelago is partitioned among the α, β_Nestedness and β_Replacement components, for pteridophytes and spermatophytes and for all endemic species and all native species. The α-diversity was more important for all native species than for endemic species and more important for pteridophytes than for spermatophytes, with the Azores showing outstanding high values of α-diversity. The β_Nestedness was higher for pteridophytes than for spermatophytes and higher for endemic species than for all native species in both pteridophytes and spermatophytes. The values of β_Replacement suggested that: (i) the spermatophyte native flora is more differentiated across islands than the pteridophyte native flora and (ii) the pteridophyte endemic flora and, especially, the spermatophyte endemic flora are more differentiated across islands than the corresponding native flora. An outstanding value of β_Replacement for endemic and all native spermatophytes was found in Hawaii, confirming the biogeographical island differentiation in this archipelago.     

Biogeographical determinants for total and endemic species richness in a continental archipelago

We examined the relationship between plant species richness and biogeographical variables (island area, island maximum elevation, distance from nearest inhabited island, distance from nearest mainland) using a data set comprising 201 islands of the Aegean archipelago. We found that endemic species richness was strongly correlated to total species richness. Single-island endemic species richness was most strongly correlated to island maximum elevation, and then to island area, with an apparent small island effect for islands smaller than 47 km². Total species richness was most strongly correlated to island area (with no apparent small island effect), and less strongly correlated to island maximum elevation. Distance from the mainland or other inhabited islands displayed limited predictive value in our data set. The slope of the relationship between species richness and geographical factors (island area, elevation, distance from island/mainland) was steeper for endemic species richness than for total richness. Finally, the different scales of endemicity (single-island endemics, island group endemics and Aegean regional endemics) displayed similar qualitative trends and only differed quantitatively. Thus, we conclude that different biogeographical factors act as drivers for total species richness than for endemic species richness.     

Effects of climate change on the distribution of plant species and plant functional strategies on the Canary Islands

Aim

Oceanic islands possess unique floras with high proportions of endemic species. Island floras are expected to be severely affected by changing climatic conditions as species on islands have limited distribution ranges and small population sizes and face the constraints of insularity to track their climatic niches. We aimed to assess how ongoing climate change affects the range sizes of oceanic island plants, identifying species of particular conservation concern.

Location

Canary Islands, Spain.

Methods

We combined species occurrence data from single-island endemic, archipelago endemic and nonendemic native plant species of the Canary Islands with data on current and future climatic conditions. Bayesian Additive Regression Trees were used to assess the effect of climate change on species distributions; 71% (n = 502 species) of the native Canary Island species had models deemed good enough. To further assess how climate change affects plant functional strategies, we collected data on woodiness and succulence.

Results

Single-island endemic species were projected to lose a greater proportion of their climatically suitable area (x ̃ = −0.36) than archipelago endemics (x ̃ = −0.28) or nonendemic native species (x ̃ = −0.26), especially on Lanzarote and Fuerteventura, which are expected to experience less annual precipitation in the future. Moreover, herbaceous single-island endemics were projected to gain less and lose more climatically suitable area than insular woody single-island endemics. By contrast, we found that succulent single-island endemics and nonendemic natives gain more and lose less climatically suitable area.

Main Conclusions

While all native species are of conservation importance, we emphasise single-island endemic species not characterised by functional strategies associated with water use efficiency. Our results are particularly critical for other oceanic island floras that are not constituted by such a vast diversity of insular woody species as the Canary Islands.     

The effects of land-use change on arthropod richness and abundance on Santa Maria Island (Azores): unmanaged plantations favour endemic beetles

We study how endemic, native and introduced arthropod species richness, abundance, diversity and community composition vary between four different habitat types (native forest, exotic forest of Cryptomeria japonica, semi-natural pasture and intensive pasture) and how arthropod richness and abundance change with increasing distance from the native forest in adjacent habitat types in Santa Maria Island, the Azores. Arthropods were sampled in four 150 m long transects in each habitat type. Arthropods were identified to species level and classified as Azorean endemic, single-island endemic (SIE), native, or introduced. The native forest had the highest values for species richness of Azorean endemics, SIEs and natives; and also had highest values of Azorean endemic diversity (Fisher’s alpha). In contrast, the intensive pasture had the lowest values for endemic and native species richness and diversity, but the highest values of total arthropod abundance and introduced species richness and diversity. Arthropod community composition was significantly different between the four habitat types. In the semi-natural pasture, the number of SIE species decreased with increasing distance from the native forest, and in the exotic forest the abundance of both Azorean endemics and SIEs decreased with increasing distance from the native forest. There is a gradient of decreasing arthropod richness and abundance from the native forest to the intensive pasture. Although this study demonstrates the important role of the native forest in arthropod conservation in the Azores, it also shows that unmanaged exotic forests have provided alternative habitat suitable for some native species of forest specialist arthropods, particularly saproxylic beetles.     

Microevolutionary patterns and processes of the Native Hawaiian colonizing fern Odontosoria chinensis (Lindsaeaceae)

Abstract.— The vascular‐plant flora of the Hawaiian Islands is characterized by one of the highest rates of species endemism in the world. Among flowering plants, approximately 89% of species are endemic, and among pteridophytes, about 76% are endemic. At the single‐island level, however, rates of species endemism vary dramatically between these two groups with 80% of angiosperms and only 6% of pteridophytes being single‐island endemics. Thus, in many groups of Hawaiian angiosperms, it is possible to link studies of phylogeny, evolution, and biogeographic history at the interspecific and interisland levels. In contrast, the low level of single‐island species endemism among Hawaiian pteridophytes makes similar interspecific and interisland studies nearly impossible. Higher levels of interisland gene flow may account for the different levels of single‐island endemism in Hawaiian pteridophytes relative to angiosperms. The primary question we addressed in the present study was: Can we infer microevolutionary patterns and processes among populations within widespread species of Hawaiian pteridophytes wherein gene flow is probably common? To address this broad question, we conducted a population genetic study of the native Hawaiian colonizing species Odontosoria chinensis. Data from allozyme analyses allowed us to infer: (1) significant genetic differentiation among populations from different islands; (2) historical patterns of dispersal between particular pairs of islands; (3) archipelago‐level patterns of dispersal and colonization; (4) founder effects among populations on the youngest island of Hawaii; and, (5) that this species primarily reproduces via outcrossing, but may possess a mixed‐mating system.     

Orchids of the West Indies: predictability of diversity and endemism

Aim We examined phytogeographical patterns of West Indian orchids, and related island area and maximum elevation with orchid species richness and endemism. We expected strong species–area relationships, but that these would differ between low and montane island groups. In so far as maximum island elevation is a surrogate for habitat diversity, we anticipated a strong relationship with maximum elevation and both species richness and endemism for montane islands. Location The West Indies. Methods Our data included 49 islands and 728 species. Islands were classified as either montane (≥ 300 m elevation) or low (< 300 m). Linear and multivariate regression analyses were run to detect relationships between either area or maximum island elevation and species richness or the number of island endemic species. Results For all 49 islands, the species–area relationship was strong, producing a z‐value of 0.47 (slope of the regression line) and explaining 46% of the variation. For 18 relatively homogeneous, low islands we found a non‐significant slope of z = −0.01 that explained only 0.1% of the variation. The 31 montane islands had a highly significant species–area relationship, with z = 0.49 and accounting for 65% of the variation. Species numbers were also strongly related to maximum island elevation. For all islands < 750 km², we found a small‐island effect, which reduced the species–area relationship to a non‐significant z = 0.16, with only 5% of the variation explained by the model. Species–area relationships for montane islands of at least 750 km² were strong and significant, but maximum elevation was the best predictor of species richness and accounted for 79% of the variation. The frequency of single‐island endemics was high (42%) but nearly all occurred on just nine montane islands (300 species). The taxonomic distribution of endemics was also skewed, suggesting that seed dispersability, while remarkable in some taxa, is very limited in others. Montane island endemics showed strong species–area and species–elevation relationships. Main conclusions Area and elevation are good predictors of orchid species diversity and endemism in the West Indies, but these associations are driven by the extraordinarily strong relationships of large, montane islands. The species richness of low islands showed no significant relationship with either variable. A small‐island effect exists, but the montane islands had a significant relationship between species diversity and maximum elevation. Thus, patterns of Caribbean orchid diversity are dependent on an interplay between area and topographic diversity.     

On Cannabis indica,Indian hemp

Background: The South Aegean Volcanic Arc (SAVA), one of the most notable geological structures of the Mediterranean Sea, is floristically well known. Nevertheless, the factors that contribute to shaping the plant species richness of the SAVA remain unclear.

Aims: To investigate the factors that affect plant species richness and identify plant diversity hotspots in the SAVA and other central Aegean islands.

Methods: We used stepwise multiple regression to test the relationship between a number of environmental factors and plant species richness in the SAVA, as well as the residuals from the species–area linear regressions of native, Greek and Cycladian endemic taxa as indicators of relative species richness.

Results: The area was confirmed to be the most powerful single explanatory variable of island species richness, while geodiversity, maximum elevation and mean annual precipitation explained a large proportion of variance for almost all the species richness measures. Anafi, Amorgos and Folegandros were found to be endemic plant diversity hotspots.

Conclusions: We have demonstrated that geodiversity is an important factor in shaping plant species diversity in the Cyclades, while mean annual precipitation, human population density and maximum elevation were significant predictors of the Greek endemics present in the Cyclades. Finally, Anafi was found to be a plant diversity hotspot in the South Aegean Sea.     

Species Richness and Patterns of Invasion in Plants, Birds, and Fishes in the United States*

We quantified broad-scale patterns of species richness and species density (mean # species/km²) for native and non-indigenous plants, birds, and fishes in the continental USA and Hawaii. We hypothesized that the species density of native and non-indigenous taxa would generally decrease in northern latitudes and higher elevations following declines in potential evapotranspiration, mean temperature, and precipitation. County data on plants (n = 3004 counties) and birds (n=3074 counties), and drainage (6 HUC) data on fishes (n = 328 drainages) showed that the densities of native and non-indigenous species were strongly positively correlated for plant species (r = 0.86, P < 0.0001), bird species (r = 0.93, P<0.0001), and fish species (r = 0.41, P<0.0001). Multiple regression models showed that the densities of native plant and bird species could be strongly predicted (adj. R² = 0.66 in both models) at county levels, but fish species densities were less predictable at drainage levels (adj. R² = 0.31, P<0.0001). Similarly, non-indigenous plant and bird species densities were strongly predictable (adj. R² = 0.84 and 0.91 respectively), but non-indigenous fish species density was less predictable (adj. R² = 0.38). County level hotspots of native and non-indigenous plants, birds, and fishes were located in low elevation areas close to the coast with high precipitation and productivity (vegetation carbon). We show that (1) native species richness can be moderately well predicted with abiotic factors; (2) human populations have tended to settle in areas rich in native species; and (3) the richness and density of non-indigenous plant, bird, and fish species can be accurately predicted from biotic and abiotic factors largely because they are positively correlated to native species densities. We conclude that while humans facilitate the initial establishment, invasions of non-indigenous species, the spread and subsequent distributions of non-indigenous species may be controlled largely by environmental factors. The U.S. Government’s right to retain a non-exclusive, royalty-free licence in and to any copyright is acknowledged.     

Comparative genetic structure and demographic history in endemic galapagos weevils

The challenge of maintaining genetic diversity within populations can be exacerbated for island endemics if they display population dynamics and behavioral attributes that expose them to genetic drift without the benefits of gene flow. We assess patterns of the genetic structure and demographic history in 27 populations of 9 species of flightless endemic Galápagos weevils from 9 of the islands and 1 winged introduced close relative. Analysis of mitochondrial DNA reveals a significant population structure and moderately variable, though demographically stable, populations for lowland endemics (F(ST) = 0.094-0.541; π: 0.014-0.042; Mismatch P = 0.003-0.026; and D((Tajima)) = -0.601 to 1.203), in contrast to signals of past contractions and expansions in highland specialists on 2 islands (Mismatch P = 0.003-0.026 and D((Tajima)) = -0.601 to 1.203). We interpret this series of variable and highly structured population groups as a system of long-established, independently founded island units, where structuring could be a signal of microallopatric differentiation due to patchy host plant distribution and poor dispersal abilities. We suggest that the severe reduction and subsequent increase of a suitably moist habitat that accompanied past climatic variation could have contributed to the observed population fluctuations in highland specialists. We propose the future exploration of hybridization between the introduced and highland endemic species on Santa Cruz, especially given the expansion of the introduced species into the highlands, the sensitivity to past climatic variation detected in highland populations, and the potentially threatened state of single-island endemics.     

Links to rare climates do not translate into distinct traits for island endemics

Current models of island biogeography treat endemic and non-endemic species as if they were functionally equivalent, focussing primarily on species richness. Thus, the functional composition of island biotas in relation to island biogeographical variables remains largely unknown. Using plant trait data (plant height, leaf area and flower length) for 895 native species in the Canary Islands, we related functional trait distinctiveness and climate rarity for endemic and non-endemic species and island ages. Endemics showed a link to climatically rare conditions that is consistent with island geological change through time. However, functional trait distinctiveness did not differ between endemics and non-endemics and remained constant with island age. Thus, there is no obvious link between trait distinctiveness and occupancy of rare climates, at least for the traits measured here, suggesting that treating endemic and non-endemic species as functionally equivalent in island biogeography is not fundamentally wrong.     

The biodiversity of the United Kingdom’s Overseas Territories: a stock take of species occurrence and assessment of key knowledge gaps

Limited financial resources for conservation and growing environmental problems make it vital to base conservation on sound scientific evidence. Small islands hold a disproportionately large amount of the worlds threatened biodiversity but it is among the least well-documented. This paper reports on the most extensive collation and synthesis of biodiversity data to date for the 14 United Kingdom Overseas Territories (UKOTs). A process of literature review and consultation produced 65,259 species records, including 32,216 native species of which 1549 were endemic to a single UKOT. The extent of knowledge of species occurrence varied both between islands and taxonomic groups. It was higher for vertebrates and vascular plants than small bodied invertebrates and non-vascular and for non-Caribbean compared to Caribbean islands, a difference that largely reflects knowledge of invertebrates. Global Red List assessments exist for 2606 species and document 111 of endemic species, 75 % of those assessed, and 291, 12 % of non-endemics, as globally threatened. Using the data to estimate true species richness suggests a further 70,000 native species, including 1800 single island endemics, remain to be documented suggesting the UKOTs as a whole may support over 100,000 native species including 3300 single island endemics.     

Endemic freshwater finfish of Asia: distribution and conservation status

Freshwater finfish species richness and level of endemism in East, and South and South‐East Asia that included 17 nations were studied using available databases, and included nation‐wise distribution, habitat types, and conservation status. The number of endemic finfish species in the region was 559, belonging to 47 families. Families Cyprinidae and Balitoridae accounted for 43.5% and 16.2% of the total number of endemic species in the region, respectively, followed by Sisoridae (25), Gobiidae (20), Melanotaeniidae (19), and Bagridae (16), and the other 41 families had at least one endemic species. Nation‐wise the most number of endemic freshwater finfish species occur in India (191), followed by China (88), Indonesia (84), and Myanmar (60). In India, the endemic species accounted for 26.4% of the native freshwater fish fauna, followed by South Korea (16.9%), the Philippines, (16.3%) and Myanmar (15.7%). Statistically significant relationships discerned between the number of indigenous and endemic species richness to land area (X_la in 10³ km²) of the nations in the region were, Y_in = 218.961 Ln(X_la) – 843.1 (R² = 0.735; P < 0.001) and Y_e = 28.445 Ln X_la?134.47 (R² = 0.534; P < 0.01), respectively, and between indigenous and endemic species richness was Y_e = 0.079X_n? 1.558 (R² = 0.235; P < 0.05). The overall conservation status of endemic finfish in Asia was satisfactory in that only 92 species were in some state of vulnerability, of which 37 species (6.6%) are endangered or critically endangered. However, the bulk of these species (83.7%) were cave‐ and or lake‐dwelling fish. However, nation‐wise, the endemic freshwater finfish fauna of the Philippines and Sri Lanka, based on the imperilment index, were found to be in a highly vulnerable state. Among river basins, the Mekong Basin had the highest number of endemic species (31.3%). The discrepancies between databases are highlighted and the need to consolidate information among databases is discussed. It is suggested that the Mekong Basin be considered as a biodiversity hotspot, and appropriate management strategies be introduced in this regard.     

Deconstructing the native–exotic richness relationship in plants

Aim Classic theory suggests that species‐rich communities should be more resistant to the establishment of exotic species than species‐poor communities. Although this theory predicts that exotic species should be less diverse in regions that contain more native species, macroecological analyses often find that the correlation between exotic and native species richness is positive rather than negative. To reconcile results with theory, we explore to what extent climatic conditions, landscape heterogeneity and anthropogenic disturbance may explain the positive relationship between native and exotic plant richness. Location Catalonia (western Mediterranean region). Methods We integrated floristic records and GIS‐based environmental measures to make spatially explicit 10‐km grid cells. We asked whether the observed positive relationship between native and exotic plant richness (R²= 0.11) resulted from the addition of several negative correlations corresponding to different environmental conditions identified with cluster analysis. Moreover, we directly quantified the importance of common causal effects with a structural equation modelling framework. Results We found no evidence that the relationship between native and exotic plant richness was negative when the comparison was made within environmentally homogeneous groups. Although there were common factors explaining both native and exotic richness, mainly associated with landscape heterogeneity and human pressure, these factors only explained 17.2% of the total correlation. Nevertheless, when the comparison was restricted to native plants associated with human‐disturbed (i.e. ruderal) ecosystems, the relationship was stronger (R²= 0.52) and the fraction explained by common factors increased substantially (58.3%). Main conclusions While our results confirm that the positive correlation between exotic and native plant richness is in part explained by common extrinsic factors, they also highlight the great importance of anthropic factors that – by reducing biotic resistance – facilitate the establishment and spread of both exotic and native plants that tolerate disturbed environments.     

Beta diversity in relation to dispersal ability for vascular plants in North America

Aim To test the hypothesis that plant species with a higher dispersal ability have a lower beta diversity. Location North America north of Mexico. Method Propagules of pteridophytes (ferns and their allies) are more vagile than propagules of spermatophytes (gymnosperms and angiosperms), and thus pteridophytes have a higher dispersal ability than do spermatophytes. The study area was divided into 71 geographical units distributed in five latitudinal zones. Species lists of pteridophytes and spermatophytes were compiled for each geographical unit. Three measures of beta diversity were used: β_sim, which is one minus the Simpson index of similarity, β_slope, which is the slope of the relationship between Simpson index and geographical distance, and β_{0.5‐distance}, which is the distance that halves the similarity from its initial value. Results Average β_sim is higher for spermatophytes than for pteridophytes, regardless of whether the data are analysed for the whole continent or for latitudinal zones. Average β_sim decreases with increasing latitude for both spermatophytes and pteridophytes. The difference in average β_sim between the two plant groups increases with increasing latitude, indicating that beta diversity decreases with increasing latitude faster for pteridophytes than for spermatophytes. When the Simpson index is regressed against geographical distance, the regression slope (β_slope) is steeper for spermatophytes than for pteridophytes, and the slope decreases with increasing latitude for both plant groups. Similarly, β_{0.5‐distance} was shorter for spermatophytes than for pteridophytes in each latitudinal zone and increased with increasing latitude for both plant groups. The results of the analyses using the three different measures of beta diversity are consistent. Main conclusions The fact that beta diversity is lower for pteridophytes with vagile propagules than for spermatophytes with less vagile propagules suggests that beta diversity is negatively related to dispersal ability.     

Are Ridge Habitats Special Sites for Endemic Plants in Tropical Montane Rain Forests? A Case Study of Pteridophytes in Ecuador

We addressed whether ridges, which are ecologically distinct from slopes, harbor specialized plant assemblages with a high representation of endemic species. We surveyed pteridophytes in 28 plots each of 400 m² in ridge and slope forests at 2,430–2,660 m at three different localities in southeastern Ecuador. Data analysis was based on those 147 species with reliable determinations and excluded 14 undetermined species. Range sizes were expressed as the latitudinal distance between the northern- and southernmost collections, and species were then assigned to range-size quartiles, with the 1st quartile including the 25% most widespread species, etc. Differences in species richness per range-size quartile were determined using G-tests and differences in abundance using ANOVAs. The recorded 147 fern species were represented by 21,800 individuals, including 106 terrestrial (7,300 individuals) and 98 epiphytic species (14,500). Ridges had fewer species than slopes, and there was no higher representation of localized species on ridges. Overall, widespread species were weakly (R²?=?0.03) but significantly more abundant than localized species. Ridges had significantly higher abundances of terrestrial – but not of epiphytic – species compared to slopes, especially among the widespread species of the 1st range-size quartile. The contribution of ridge habitats to overall pteridophyte diversity and as habitats for endemics in our study region is low. Methodologically, the separation of species into range-size classes in an ecological study is novel and effective, as statistically significant patterns were found only for species belonging to the 1st or 4th quartiles.     

Morphology,ultrastructure and phylogenetic affinities of the single-island endemic Anthoceros cristatus Steph. (Ascension Island)

Oceanic islands, due to their geographical isolation, number, precisely defined boundaries and their geomorphological and climatic diversity, have provided enormous insights into speciation, dispersal, adaptive radiations and macroecological processes. One of the key components of these island studies is the role of single-island endemics (SIEs) as, in many instances, island biogeography models use the proportion of SIEs to infer evolutionary processes. It is, therefore, imperative to undertake critical taxonomic revisions to evaluate SIEs because changes in the number of SIEs have a key impact on downstream biogeographic analyses. We revise the special case of a putative SIE Anthoceros cristatus on Ascension Island using light and electron microscopy, as well phylogenomic tools. A. cristatus lies within the A. agrestis/A. punctatus complex but differs from the sister species A. agrestis and A. punctatus in spore morphology and gametophytic lamellae fringed with caducous marginal cells. The present confirmation, from both molecules and morphology, of the SIE status of Anthoceros cristatus and its restricted distribution on the Island makes the preservation of its habitat a conservation priority. Ascension Island is the tip of an undersea volcano that is thought to have emerged from the ocean 1 million years ago with an area of approximately 91?km², with Green Mountain as the highest elevation (～859?m a.s.l.). Ascension has a relative low bryophyte species diversity of 87 spp but this includes 12 endemics (～14%); a much higher level of endemism than on the far more speciose Macaronesian Islands.     

Tracing ancestry with chromosomal sequences

Most of the large Drosophila species of Hawaii are single-island endemics. Chromosomal sequences show that species at the new end of the archipelago have been derived stepwise from ancestral populations on older islands. The oldest high island has an endemic species with sequences that match some in the Nearctic-Palearctic robusta species group. Colonization from a continent by long-distance dispersal seems a likely origin for the Hawaiian drosophilids. Telmatogeton, a worldwide genus of marine midges, has five Hawaiian species inhabiting freshwater streams. Chromosomal sequences of a marine species in Hawaiian waters match the freshwater forms, indicating colonization from the ocean.     

Evaluating dominance as a component of non-native species invasions

Many studies have quantified plant invasions by determining patterns of non‐native species establishment (i.e. richness and absolute cover). Until recently, dominance has been largely overlooked as a significant component of invasion. Therefore, we re‐examined a 6‐year data set of 323 0.1 ha plots within 18 vegetation types collected in the Grand Staircase‐Escalante National Monument from 1998 to 2003, including dominance (i.e. relative cover) in our analyses. We specifically focused on the non‐native species Bromus tectorum, a notable dominant annual grass in this system. We found that non‐native species establishment and dominance are both occurring in species‐rich, mesic vegetation types. Therefore, non‐native species dominance may result despite many equally abundant native species rather than a dominant few, and competitive exclusion does not seem to be a primary control on either non‐native species establishment or dominance in this study. Unlike patterns observed for non‐native species establishment, relative non‐native species cover could not be predicted by native species richness across vegetation types (R² < 0.001; P = 0.45). However, non‐native species richness was found to be positively correlated with relative non‐native species cover and relative B. tectorum cover (R² = 0.46, P < 0.01; R² = 0.17, P < 0.01). Analyses within vegetation types revealed predominantly positive relationships among these variables for the correlations that were significant. Regression tree analyses across vegetation types that included additional biotic and abiotic variables were a little better at predicting non‐native species dominance (PRE = 0.49) and B. tectorum dominance (PRE = 0.39) than at predicting establishment. Land managers will need to set priorities for control efforts on the more productive, species‐rich vegetation types that appear to be susceptible to both components of invasion.     

The distribution of native and exotic plants in a naturally fragmented sagebrush-steppe landscape

We tested two general hypotheses for the diversity of native and exotic plants in an undisturbed, naturally fragmented sagebrush-steppe landscape in SE Idaho, USA, evaluating whether the MacArthur–Wilson hypothesis of island biogeography or a suite of environmental variables explained the distributions of native and exotic plants. We also tested a third hypothesis, which incorporated assumptions about the origin of exotic plants and their interaction with native plants. Of the three hypotheses we tested, the hypothesis that included exotic species best explained the diversity of the native plant community. The MacArthur–Wilson model of island biogeography did not explain the diversity of native (R ² = 0.13) or exotic plants well (R ² = 0.11), and the model fit the data poorly. A model of environmental variables better explained the diversity of native (R ² = 0.48) and exotic plants (R ² = 0.57), but it also fit the data poorly. Instead, proximity to a railroad explained the cover (R ² = 0.59) and richness of exotic plants (R ² = 0.63), which then explained the species richness of native plants (R ² = 0.34), and the model fit was adequate and had the lowest AIC value. This suggests that the transportation corridor had a significant, though indirect, effect on the native plant community, even in this undisturbed area. Moreover, explained variance, model fit, and the AIC model selection criteria favored the model with the railroad and exotic species over the M–W and environmental models. Since the habitat patches we studied were largely undisturbed by people and their activities, our results further suggest that the transportation corridor influenced the distribution of exotic plants by serving as a vector for colonization, rather than as a source of disturbance. Additionally, the results suggest that exotic plant species have had a negative effect on the diversity of the native plant community and have changed its composition. The results also support the inference that the nascent exotic plant community is influenced by source-sink (Pulliam in Am Nat 132:652–661, 1988) and assembly dynamics. In contrast, the native plant community appears to be more strongly influenced by environmental conditions associated with an elevational gradient, but there is evidence that the native community also has undergone directional change in species composition, associated with the invasion by non-native species.